Biomed 

UK 

520 

F398 

1900 


0 
8  I 

7 

4 


UNIVERSITY  OF  CALIFORNIA 
AT  LOS  ANGELES 


FERNWORT    PAPERS, 

PRESENTED  AT  A  MEETING 

OF 
FERN  STUDENTS, 

HELD  IN   NEW  YORK  CITY  JUNE  27,  1900, 

UNDER   THE  AUSPICES  OF  THE 

LINN^AN   FERN  CHAPTER. 


Issued  December  20,  1900. 


Printed  for  the  -Linnaean  Fern  Chapter. 


BlNGHAMTON,   N.   Y. 

WIZARD  N.  CI<UTE  &  CO  : 
1900. 


THE  LINN^AN  FERN  CHAPTER. 
OFFICERS  FOR  1901. 

President 
WILLIAM  R.  MAXON,          U.  S.  National  Museum, 

Washington,  D.  C. 

Vice-Presiden  t 
B.  D.  GILBERT,  Clayville,  N.  Y. 

Secretary 
Miss  MARGARET  SLOSSON,  Andover,  Mass. 

Treasurer 
JAMES  A.  GRAVES,  Susquehanna,  Pa. 


This  Chapter,  founded  in  1893,  has  as  its  aim 
the  study  of  ferns  by  correspondence,  and  the  publi- 
cation of  the  results  of  such  study.  It  comprises 
nearly  125  members  from  all  parts  of  the  United 
States,  and  from  Canada  and  Great  Britain.  All  who 
are  interested  in  ferns  are  cordially  invited  to  join. 
Dues  are  one  dollar  annually  ;  the  initiation  fee  of 
one  dollar  includes  the  dues  of  the  first  year.  Mem- 
bers receive  without  cost  the  Fern  Bulletin  quar- 
terly and  Annual  Reports ;  other  publications  either 
without  cost  or  at  a  rate  considerably  lower  than 
available  to  other  than  Chapter  members.  For  full 
information  address  either  the  President  or  Secretary. 

FOR  PRICE  LIST  OF  PUBLICATIONS,  SEE  FOURTH  PAGE  OF  COVER. 


^f^^e^ 

O  , 

^  /  \/w  n  »  2 


FERNWORT    PAPERS, 

PRESKNTED  AT  A  SECOND  MEETING  OF  FERN  STUDENTS,  HELD 
IN  NEW  YORK  CITY,  JUNE  27,  1900,  UNDER 

THE    AUSPICES   OF   THE    LlNNAEAN    FERN    CHAPTER.* 


•  the  Chapter.  Issued  December  20.  1900 


THE  GENUS  ISOETES  IN  NEW  ENGLAND. 

Bv  A.  A.  EATON. 


In  the  three  seasons  that  have  elapsed  since  the  appearance  of 
Dodge's  "Ferns  and  Fern  Allies  of  New  England,"  the  study  of 
our  local  species  of  the  genus  hoetes  has  progressed  so  far  that  it 
seems  advisable  at  this  time  to  present  some  of  the  results,  and 
state  some  of  the  problems,  in  order  to  elicit  the  co-operation  of 
collectors  in  clearing  up  some  of  the  unsettled  questions.  As  the 
settling  of  these  questions  will  pave  the  way  to  a  larger  and  more 
comprehensive  work,  this  paper  may  be  considered  as  preliminary 
only . 

It  appears  advisable  to  give  a  history  of  the  genus,  so  far  as  our 
chosen  territory  is  concerned,  and  in  doing  this  we  can  do  no  better 
than  to  adapt  and  extend  Engelmann's  chronological  history  in  his 
comprehensive  account  of  the  genus,  given  in  Trans.  St.  Louis 
Acad.  4:358-390.  1882. 

According  to  this,  the  first  collection  was  at  Upbridge,  Mass., 
by  Robbins,  in  1831.  It  was  referred  to  riparia. 

1840.     Robbins  collected  lacustris  in  the  same  town. 

1843.     Tuckerman  collected  lacustris  in  Echo  Lake,  N.  H. 

1845.     Robbins  found  echinospora  Braunii  in  Massachusetts. 

1848.  Tuckerman  found  near  Boston  the  species  which  bears 
his  name. 

1856.  Engelmann  found  echinospora  Braunii  in  Lake  Winni- 
piseogee,  New  Hampshire. 

1857.  E.  D.  Eaton  found  Engelmanni  in  New  England. 


*  For  an  account  of  this  Meeting,  see  Fern  Bulletin  for  July,  1900.  The  first 
gathering  of  those  interested  in  American  fern  study  occurred  in  Boston, 
August  24,  1898,  and  the  papers  read  were  published  early  in  1899,  under  the 
title,  "  Papers  Presented  at  the  Boston  Meeting." 


200775 


1    1860.     Boott  discovered  echinospora  muricata. 

1865.     Boott  rediscovered  Tuckermani. 

1867.     Boott  collected  echinospora  Boottii. 

1878.  Pringle  collected  echinospora  robusta  in  Lake  Cham- 
plain. 

Engelmann  also  mentions  other  New  England  collections,  but 
they  are  not  given  in  his  list.  After  his  time  there  is  a  hiatus  in 
publications,  but  collectors  were  accumulating  undigested  material, 
and  wrongly  referring  it  often.  The  localities  for  Braunii  have  be- 
come so  numerous  that  it  appears  useless  to  enumerate  them. 
They  occur  in  all  of  the  New  England  States. 

1887.  Kennedy  found  what  is  now  known  as  Tuckermani 
borealis  in  Somes  stream,  Mt.  Desert. 

1889.  Dr.  C.  B.  Graves  found  echinospora  muricata  in  Con- 
necticut. 

1892.  Fernald  collected    Tuckermani   and   Harveyi    at    Mt. 
Desert,  and  Rand  discovered  what  is  now  the  type  of  heterospora 
at  the  same  place. 

1893.  Pringle  and  Fernald  collected  the  types  of  hieroglyphica 
at  St.  Francis  lakes.     Dodge  discovered  Engelmanni  at  Newbury. 

1894.  Dodge  discovered    Tuckermani  and  Braunii  at  New- 
buryport. 

1895.  Coville  discovered  Tuckermani  borealis  at  Kennebago 
lakes,  and  hieroglyphica  at  the  Rangeleys.     Raynal  Dodge  and 
myself  found    Tuckermani  at  Essex,  Mass.,  and  severally  discov- 
ered it  at  Amesbury.     I  also  found  it  at  Kingston,  N.  H.,  and  dis- 
covered muricata  at  Kingston  and  East  Kingston,  where  I  found 
the  types  of  Eatoni  and  one  specimen  of  Dodgei  in  a  pond  running 
into  Kingston.     Graves  collected  Gravesii  in  Connecticut. 

1896.  I  discovered  Dodgei  at  Kingston  in  abundance,  foveo- 
lata,  Eatoni,  echinospora   robusta,  muricata   and    Tuckermani  at 
Epping,  N.  H.,  and  the  last  at  Nottingham.     Dr.  Graves  collected 
Tuckermani  in  southern  Connecticut. 

During  the  years  1895  and  1896  I  found  Engelmanni  to  be  com- 
mon in  Rockingham  county,  N.  H.,  and  echinospora  Braunii  in 
many  places. 

1898.  Harvey  discovered  the  type  of  Harveyi  at  Pushaw  pond, 
Oldtown,  Maine.     I  found  Eatoni  at  Amesbury,  and  Dodge  at  New- 
bury, Mass. 

1899.  Harvey  found  Tuckermani  botealis  at  Oldtown,  Maine, 
hieroglyphica  and  muricata  at  Moosehead  lake.     1  traced  Eatoni, 


010924 


— 3— 

foveolata,  and  muricata  to  Newmarket,  N.  H.  Underwood  found 
some  peculiar  plants,  not  clearly  referable  to  any  published  species, 
at  Goshen,  Conn.  The  spores  are  either  abortive  or  unripe,  and 
unsatisfactory  for  study."" 

It  is  hoped  collectors  will  call  my  attention  to  the  many  gaps  in 
this  list,  that  I  may  fill  them. 

A  critical  study,  however,  shows  the  remarkable  fact  that  ri- 
paria  and  lacustris,  tho  formerly  considered  common,  are  very  rare 
and  have  not  been  collected  in  recent  years.  It  is  true  they  have 
been  reported  from  various  localities,  but  all  such  specimens  which 
have  been  accessible  to  me,  have  proven  to  be  either  Engelmanni, 
Tuckermani,  Tuckermani  borealis,  or  an  undescribed  species. 
It  is  unsafe  to  draw  conclusions  until  some  of  the  old  material  has 
been  re-examined,  which  I  hope  will  be  in  the  near  future  ;  but 
there  is  a  growing  probability  that  the  former  has  rarely  been  met 
north  of  the  Delaware,  and  that  the  latter  in  its  typical  state  is  not 
American.  I  cannot  agree  with  Dodge  in  his  determinations  as 
regards  these  species,  and  identify  them  mostly  as  Tuckermani. 
Incidentally,  his  saccharata  from  the  Merrimac  is  immature  Braunii. 

There  is  one  other  point  in  his  work  to  which  I  take  exception, 
and  that  is  the  theory  of  hybridity  of  our  species.  Tho  this  may 
well  occur,  I  have  never  seen  a  case  which  I  thought  called  for  that 
explanation.  Eatoni,  as  is  well  known,  usually  bears  only  female 
sporangia,  only  one  plant  in  two  or  three  hundred  bearing  male 
spores,  and  then  the  sporanges  occupy  no  regular  zone,  but  are 
intermixed  with  the  others.  It  is  evident,  then,  that  this  would  be 
a  very  good  plant  to  experiment  upon,  as  the  macrospores  could  be 
easily  obtained  without  microspores.  This  has  been  done  by  Mr. 
T.  C.  Palmer,  who,  I  hope,  will  soon  publish  the  results.  I  will 
anticipate  this,  however,  by  stating  that  while  straight  cultures 
yielded  plants,  attempted  crosses  between  Eatoni  and  Dodgei  or 
Engelmanni  were  negative.  So  we  have  presumptive  proof  that 
hybridity  is  extremely  rare,  if  not  altogether  absent.  Mr.  Dodge 
based  his  presumption  on  spore  characters  only,  a  very  unsatisfac- 
tory character,  inasmuch  as  the  sculpture  of  spores  of  all  species  is 
very  variable,  in  the  echinosporas  often  confluent  into  walls,  espe- 
cially in  muricata;  while  in  Tuckermani  spores  are  often  found 
mixed  among  normal  ones  that  have  the  crests,  or  some  of  them, 
broken  up  into  spinules.  As  these  spores  have  been  found  in 

*  Additional  specimens  from  I/ynn,  Conn.,  collected  by  Dr.  Graves,  prove 
this  to  be  an  undescribed  species.    It  is  inserted  under  the  name  of  /.  Gravssii. 


sporanges  with  normal  ones,  I  think  them  explainable  better  on  the 
theory  of  accidental  variation,  than  on  that  of  hybridity,  especially 
as  they  are  found  where  the  species  do  not  grew  in  juxtaposition. 
My  investigations  on  this  point  have  been  extensive,  covering 
thousands  of  plants,  and  my  conclusion  is  that  hybridity  in  this 
genus,  though  not  impossible,  nor  even  improbable  so  far  as  ex- 
ternals go,  is  at  least  extremely  rare  among  New  England  species. 

I  wish  here  to  pay  tribute  to  the  general  excellence  of  Mr. 
Dodge's  treatment  of  the  genus,  a  more  practical  account  never 
having  been  written  ;  and  for  the  beginner,  I  can  conceive  of  no  bet- 
ter directions  than  are  contained  in  his  notes.  I  wish  I  could  say 
as  much  for  Britton  and  Brown's  more  pretentious  "Illustrated 
Flora,"  but  to  anyone  with  more  than  the  most  superficial  knowl- 
edge of  the  genus,  their  treatment  is  very  disappointing. 

Considerable  labor  has  been  expended  by  Durieu,  Braun,  and 
Engelmann,  to  devise  some  system  of  classification  that  would  hold 
water,  but  with  ill  success,  as  they  themselves  testify  ;  for  while  it 
is  comparatively  easy  to  classify  species  of  a  limited  area,  the  sys- 
tem fails  when  applied  to  the  genus  as  a  whole.  Perhaps  the  most 
natural  grouping  is  into  the  three  divisions  Aquaticae  or  Submersae, 
Amphibiae,  and  Terrestres,  and  if  applied  with  sufficient  looseness 
will  hold.  But  Submersae  in  Europe  are  bilobed  and  without 
stomata,  consisting  of  the  species  lacustris  and  echinospora.  When 
applied  to  the  genus  as  a  whole,  however,  we  find  Gunnii  and  elatior 
of  Tasmania  three  lobed,  and  also  find  our  varieties  of  echinospora 
with  stomata,  thus  placing  them  in  a  different  group  from  the  type, 
as  indeed  they  appear  from  habit  to  fall.  The  system  fails,  too, 
when  taken  in  its  original  sense  in  the  division  Amphibiae.  All 
Old  World  species  of  this  group  are  three  lobed,  and  this  was  made 
use  of  in  the  classification  ;  but  not  one  of  our  numerous  species  of 
eastern  America  is  three  lobed,  and  only  four  from  North  America, 
Nuttallii,  Cubana,  Orcuttii  and  minima,  are  so  constituted.  The 
only  works  treating  of  the  genus  as  a  whole,  do  not  tend  to  clear 
up  the  matter.  Baker,  in  "Fern  Allies,"  treats  the  genus  very 
superficially,  while  Motelay's  more  pretentious  work  is  full  of  inac- 
curacies, nearly  two  pages  being  given  to  "  errata,"  and  these  cov- 
ering a  small  percentage  of  those  contained  in  the  work.  Though 
a  help  in  many  ways,  the  critical  student  hesitates  to  place  much 
weight  on  statements  accompanied  by  such  errors. 

The  group  Terrestres  is,  it  seems  to  me,  unnecessarily  limited 
to  the  two  species  with  persistent  leaf  bases.  Nuttallii,  fiutleri,  and 


— 5— 

Montezumas  of  America,  placed  beside  Duriaei  or  hystrix  of 
Europe,  could  not  be  separated  by  superficial  examination,  this  one 
point  aside  ;  and  some  forms  of  Duriaei  and  hystrix,  those  growing 
in  damp  places,  habitually  have  no  scales.  On  the  other  hand,  the 
three  species  mentioned  can  be  separated  at  a  glance  from  melano- 
poda,  Mexicana,  or  Pringlei.  Leaf  sections  also  are  identical  in 
the  Terrestres,  the  leaves  being  short,  setaceous,  with  very  small 
cavities,  and  correspondingly  wide  dissepiments,  and  with  four 
stout  bast-bundles  (three  in  Nuttallii},  while  the  others  named,  as 
well  as  most  European,  African,  and  Asian  species  which  I  have 
examined,  have  stout  leaves,  four  primary,  and  several  accessory 
bast-bundles,  narrower  dissepiments,  and  correspondingly  larger 
cavities.  I  leave  a  further  discussion  of  these  points  for  a  future 
occasion,  especially  as  they  bear  only  remotely  on  the  immediate 
subject  in  hand.  I  would  remark,  however,  that  Engelmann  is  at 
variance,  and  I  think  rightly  so,  with  European  authors  in  his  con- 
ception of  the  subdivisions. 

Probably  the  main  reason  that  our  species  of  Isoetes  are  not 
better  known,  both  in  habit  and  distribution,  is  the  difficulty  novices 
experience  in  finding  them.  Their  unattractive  appearance  and  the 
tedious  process  of  analysis  by  use  of  a  compound  microscope,  re- 
quired to  determine  them  with  accuracy,  have  also  conspired  to 
prevent  general  study.  No  character  given  can  be  expected  to  be 
true  in  all  phases.  The  leaves  are  long  or  short,  according  to  the 
stage  of  the  water  ;  they  are  stout  or  slender,  and  the  trunk  large 
or  small,  to  correspond  to  the  general  vigor  of  the  plant.  The 
outer  sporanges  are  usually  nearly  round,  while  the  inner  are  two  to 
four  times  as  long  as  broad.  The  velum  varies  greatly  in  some 
species,  in  certain  forms  of  echinospora  varying  from  ^-%  indusi- 
ate,  and  I  have  seen  specimens  from  Maine  and  Amesbury,  Mass. , 
where  a  pin  head  would  cover  the  opening.  I  should  have  consid- 
ered this  a  good  variety,  had  not  fresh  specimens  from  the  same 
place  possessed  a  normal  velum.  The  spores  vary  greatly  in  size, 
and  one  may  always  expect  to  find  some  larger  than  the  descrip- 
tion allows.  Location  appears  to  regulate  this.  Juckermani,  for 
instance,  is  normal  in  this  respect  in  most  Massachusetts  localities, 
while  New  Hampshire  and  Maine  plants  usually  measure  10-20  // 
more. 

I  know  of  no  character  more  unreliable  than  the  sculpture  of 
the  spores.  It  is  always  easy  to  recognize  Engelmanni  in  New 
England,  but  southward  even  this  breaks  up  into  several  varieties. 


Eatoni  usually  has  its  peculiar  convoluted  ridges,  but  one  often  finds 
a  spore  which  is  cristate.  Dodgei  shows  the  same  tendency  to 
vary,  and  to  a  greater  extent  Tuckermani,foveolata,  and  the  echino- 
sporas.  In  the  latter  one  may  at  times  find  a  spore,  or  perhaps  sev- 
eral in  a  sporangium,  with  crests  running  across  the  upper  faces, 
while  even  the  spines  below  are  variable.  Other  spores  grown 
with  them,  will  have  very  long,  slender,  forked  spines.  Indeed,  I 
do  not  at  present  feel  competent  to  fix  the  limits  of  the  varieties, 
but  leave  them  where  Engelmann  did  ;  or,  where  specimens  have 
been  referred  to  one  or  another  of  the  varieties,  the  arrangement  is 
tentative  only. 

I  would  remark,  however,  that  while  most  specimens  from  my 
neighborhood  answer  well  to  the  descriptions  of  muricata  and 
Boottii,  northern  and  mountain  specimens  usually  have  scattered, 
short,  blunt,  flat  spinules,  most  nearly  corresponding  to  Braunii, 
which,  as  originally  described,  is  a  mountain  form. 

The  leaves  of  all  species  bear  a  central  bast-bundle,  but  none  of 
ours  have  more  than  four  peripheral  ones,  and  only  four  have  them 
at  all,  Engelmanni,  Eatoni,  Dodgei  and  Gravesii.  In  well  devel- 
oped plants  of  the  first,  all  four  are  pretty  constant,  but  young  or 
starved  plants  are  liable  to  have  only  one  or  two,  perhaps  none. 
Eatoni  is  oftener  without  than  with  them. 

All  our  species  habitually  have  bi-lobed  trunks,  but  tri-lobed 
ones  are  not  infrequently  found,  and  in  one  species,  Tuckerwani, 
I  find  about  25  per  cent,  tri-lobed,  and  have  found  one  four-  and 
another  seven-lobed.  While  these  plants  appear  to  have  been 
chiefly  tri-lobed  from  the  seedling,  occasionally  one  is  found  which 
is  apparently  in  process  of  transformation,  One  notable  case  was 
a  plant  oifoveolata,  collected  at  Newmarket,  N.  H.,  in  1899,  which 
bore  a  distinct  lesion  on  one  side,  in  which  roots  were  forming. 
The  cortex  between  the  lobe  and  lesion  had  not  yet  given  way,  but 
in  the  natural  course  of  events  would  soon  have  decayed,  and  prob- 
ably the  root-bearing  area  would  have  persisted,  the  plant  thus  be- 
coming tri-lobed.  Indeed,  in  the  four-  and  seven-lobed  Tucker- 
inani  specimens,  there  was  evidence  that  a  similar  accident  had  be- 
fallen them,  and  we  might  conjecture  from  this  that  the  lower  part 
of  the  trunk  is  capable  of  becoming  root-bearing,  provided  the  cor- 
tex be  ruptured.  Nothing  conclusive  has  yet  been  observed  on  this 
point  and  it  needs  a  series  of  experiments  to  demonstrate  the  truth  • 
or  falsity  of  the  hypothesis. 


In  the  following  arrangement  I  have  based  the  key  on  charac- 
ters which  will  most  easily  allow  the  determination  of  species  with- 
out recourse  to  the  compound  microscope.  Though  not  perfect,  I 
hope  it  will  be  helpful  to  the  general  student, 

SYSTEMATIC  ARRANGEMENT. 
Group  i. 

Plants  habitually  under  water  in  the  driest  seasons,  growing  in 
sand  or  gravel.  Bast  absent,  and  stomata  few  or  none. 

Submersae. 

Leaves  1.5  to  2mm.  in  diameter,  stiffly  erect,  stomata  none. 

1015  cm.  long,  spores  (averaging  under  600  //)  covered  with 
short  crests,  rarely  a  little  reticulate  below.  i.  lacustris. 

5-8  cm.  long,  spores  larger  (averaging  over  600  n,  at  times 
over  1000  /*),  of  various  shapes,  more  densely  cristate, 
crests  anastamosing  but  not  reticulated.  2.  heterospora . 

Leaves  spiral  or  recurved,  stomata  few  or  none. 

Leaves  very  slender,  under  i  mm.  in  diameter,  reddish  when 
young,  becoming  olive-green,  8-15  cm.  long,  stomata  in 
single  series  over  air  cavities,  or  none,  macrospores  wavy 
crested  above,  more  or  less  reticulate  below,  600  /j.  or  less 
in  diameter.  3.  Tuckerniani. 

Leaves  dull  green,  stouter,  about  imm.,  not  spiral,  slightly 
recurved,  stomata  not  seen,  spores  larger,  averaging  above 
650  /*.  33.  Tuckermani  borealis. 

Leaves  stouter,  over  i  mm.  in  diameter,  shorter,  8-iocm., 
rigidly  recurved,  spores  less  than  600  fj.,  loosely  covered 
with  vermiform  wrinkles.  4.  hieroglyphica. 

Leaves  very  stout,  2.5-3mm.,  short,  5-6cm.,  purple  bronze  in 
color,  rigid,  recurved,  spores  as  in  No.  3.  5.  Harveyi. 

Group  2. 

Plants  growing  usually  in  mud  on  borders  of  ponds  or  rivers, 
inundated  most  of  the  year,  but  fruiting  as  the  water  recedes.  Sto- 
mata abundant  but  peripheral  bast-bundles  none.  Never  far  from 
water,  and  always  in  very  damp  soil.  Amphibiae. 

Spores  reticulated  below,  jagged  cristate  above,  much  as  in 
No,  3,  but  leaves  stouter  and  erect.  6.  riparia. 

Spores  smaller,  averaging  440  /*,  covered  with  very  small 
pits.  7-  foveolata. 

Spores  covered  with  spinules,  stomata  absent. 

8.  echinospora,  not  American. 


Stomata  present. 

Mo.st  of  the  spines  short,  broad,  usually  retuse,  leaves  25  or 
less.  8a.  echinospora  Braunii. 

Like  No.  8a,  but  leaves  25-70,  and  more  densely  stomatose. 
8b.  echinospora  robusta. 

Spines  long  and  slender,  sharp  or  forked,  leaves  stiffly  erect. 

Sc.  echinospora  Boottii. 

Spines  often  mixed  with  short  crests,  otherwise  as  in  No.  8a, 
spores  larger,  leaves  long  and  slender. 

8d.  echinospora  muricata. 

Group  3. 

Plants  growing  in  ditches  or  near  the  borders  of  ponds  at  higher 
levels  than  the  preceding,  emersed  during  the  greater  part  of  the 
summer.  Stomata  many,  and  bast-bundles  (in  our  species)  four. 

Palustres. 
Spores  smaller  (450  ^  or  less). 

Closely  set  with  irregular,  thick,  rough,  anastamosing 

ridges.  9.  Eatoni. 

Densely  covered  with  short,  truncate  columns.    10.    Gravesii 

Spores  larger  (averaging  over  550  ft)  with  irregular,  usually  scat- 
tered, variously  anastamosing  crests,  ii.  Dodgei. 

Spores  medium  ( averaging  about  450  fi } ,  regularly  honeycomb- 
reticulated. 
Plants  larger,  with  stouter  leaves,  and  four  bast-bundles, 

12.  Engehnanni. 

Plants  weaker,  leaves  15  or  less,  often  without  bast-bundles. 
I2a.   Engehnanni  gracilis. 

I.       I.   LACUSTRIS     L. 

Quite  common  in  clear  waters  in  northern  Europe.  Variously 
reported  from  North  America,  but  most  reports  have  proven  erro- 
neous. The  nearest  approach  to  it  I  have  yet  met,  was  collected 
by  Coville  in  Kennebago  lake,  Maine,  in  1895.  I  have  referred  his 
plants  doubtfully  to  Tuckermani  borealis. 

2.  I.  heterospora    n.  sp. 

Trunk  bi-lobed,  leaves  50-75,  5-8cm.  long,  very  stiffly  erect, 
nearly  2mm.  in  diameter,  tapering  to  a  sharp  point,  wanting  stomata 
or  bast :  velum  J/j-%  indusiate  :  sporangium  spotted,  often  thickly 
so,  with  dark  cells  ;  macrospores  normally  540-675  //,  but  occasion- 
ally i  loo  or  even  1134  n,  densely  covered  with  thick,  jagged,  con- 
voluted crests,  often  honeycomb-reticulated  below  ;  microspores 
30.8-39.6  ft,  averaging  35x27  p,  dark  brown,  papillose. 


This  species  differs  from  locus fris in  its  shorter,  more  numerous, 
tapering  leaves,  spotted  sporangium,  papillose  microspores,  and 
larger  macrospores. 

It  is  remarkable  for  the  shape  and  size  of  the  spores.  Nearly 
all  the  sporanges  of  my  plants  are  unripe,  but  good  spores  are  found 
among  the  roots.  In  these  the  sculpture  opens  into  a  network. 
Normal  spores  are  found,  but  most  of  them  bear  evidence  of  having 
grown  singly  or  in  pairs  in  the  mother  cells,  as  does  Selaginella  ru- 
pestris.  These  are  spherical  or  hemispherical,  without  commisures 
and  with  no  equatorial  belt,  or  with  it  misplaced,  often  inclosing  a 
very  small  space  at  one  end.  The  microspores  also  vary,  some- 
times reaching  44  p. 

Deer  brook  beach,  Jordan  Pond,  Mt.  Desert  Island,  Maine, 
Rand.  Quoted  in  Redfield  and  Rand's  Flora  as  Braunii.  South 
shore  of  Jordan  Pond,  September  10,  1894,  Rand. 

Type  in  the  A.  A.  Eaton  herbarium. 

3.     I.  TUCKERMANI   A.  Br. 

Formerly  considered  rare,  this  species  appears  to  be  quite  com- 
mon in  New  England,  and  any  large  pond  with  sandy  shores  may 
be  expected  to  yield  it,  especially  if  a  little  silt  has  been  deposited 
on  the  sand.  It  is  usually  found  in  "submersed  pastures,"  the  belt 
where  littoral  vegetation  extends  beneath  the  surface  of  the  water 
for  a  short  distance  after  the  lowest  stage  of  water  has  been  reached. 
I  quite  accidentally  discovered  in  1895  that  this  species  was  tri-lobed 
in  many  instances,  and  some  localities  yield  25  per  cent,  of  tri-lobed 
plants.  Baker  (Fern  Allies,  p.  126)  appears  to  be  the  first  to  men- 
tion stomata  in  this  species,  though  the  fact  was  discovered  by  Mr. 
Dodge  and  myself  quite  independently.  Plants  taken  from  inun- 
dated situations  usually  have  none,  but  when  growing  on  the  bor- 
ders of  ponds,  a  few  leaves  may  usually  be  found  which  show  them. 

Distinguishable  at  sight  by  its  very  slender,  spiral  or  recurved, 
reddish  leaves.  Spores  of  northern  specimens  are  much  larger  than 
those  from  central  Massachusetts,  so  far  as  seen. 

At  Pautuckaway  Pond,  Nottingham,  N.  H.,  and  Kimball's 
Pond,  Amesbury,  Mass.,  I  find  a  few  plants  which  in  the  field  have 
been  taken  for  this  species,  but  the  leaves  are  larger,  the  sporangia 
dark  spotted,  as  much  so  as  most  specimens  of  Howellii.  It  ap- 
pears to  be  a  new  species,  but  material  is  inadequate  for  descrip- 
tion. 

Mt.  Desert,  Rand,  Fernald:  Oldtown,  Maine,  Harvey.     Com- 


mon  in  Rockingham  county,  N.  H.,  and  known  from  several  ponds 
in  Essex  county,  and  near  Boston,  Mass.,  ;  also  from  North  Ston- 
ington,  and  Ledyard,  Conn.,  Graves. 

3a.     I.  Tuckermani  borealis  n.  var. 

Trunk  bi-lobed,  leaves  1020,  3-8cm.  long,  imm.  thick,  slightly 
recurved  at  tip  :  sporanges  sometimes  showing  a  few  spots  ;  mac- 
rospores  600-783  /u,  sculptured  as  in  the  species,  but  the  markings 
larger. 

This  might  perhaps  as  well  be  considered  a  variety  of  lacustris, 
but  the  smaller  leaves,  wider  velum,  spotted  sporanges  and  reticu- 
lated spores,  are  characters  enough  to  separate  it,  even  if  that  spe- 
cies were  not  so  extremely  rare. 

Somes  stream,  Mt.  Desert,  August  13,  1887,  Kennedy;  pond 
north  of  Long  Pond,  Mt.  Desert,  September  22,  1892  (with  Harveyi 
and  Tuckermani }  Fernald;  Somes  stream,  September  30,  1893, 
Rand;  Kennebago  lake  (leaves  only  2.5-5011.  long),  January  12, 
1895,  Coville,  no.  78  ;  Somes  stream,  Mt.  Desert,  September  14, 
1895,  Rand;  Pushaw  Pond,  Oldtown,  Maine,  August  21,  1899, 
Hervey.  I  also  occasionally  find  it  in  Lamprey  river,  at  Epping, 
New  Hampshire. 

Type  in  the  herbarium  of  A.  A.  Eaton  ;  co-types  in  the  U.  S. 
National,  and  University  of  Minnesota  herbaria. 

4.     I.  hieroglyphica  n.  sp. 

Aspect  of  Tuckermani :  trunk  bi-lobed  :  leaves  10-20,  6-7. 5011. 
long,  recurved,  i-2mm.  in  diameter,  blunt  at  tip :  velum  ^  indusi- 
ate,  sporangium  unspotted  ;  macrospores  486-590  /a,  polished,  cov- 
ered with  vermiform,  subconfluent  and  somewhat  reticulated  ridges, 
becoming  naked  next  the  equator  ;  microspores  31-44  v,  averaging 
39  p,  distinctly  verrucose. 

The  spores  are  unique  in  appearance  ;  the  ridges  are  very  bold, 
and  the  rest  of  the  surface  unmarked.  I  have  seen  nothing  like  it 
from  any  quarter,  but  Motelay's  illustration  oiflaccida  spores  gives 
a  fair  idea  of  their  appearance,  though  utterly  unlike  the  species  he 
intends  to  represent. 

St.  Francis  lakes,  Maine,  type,  Pringle;  also  Fernald  from  the 
same  place  ;  Moosehead  lake,  Harvey;  Rangeley  lakes,  Coville. 

Type  in  the  herbarium  of  A.  A.  Eaton  ;  co-types  in  the  U.  S. 
National,  and  the  University  of  Minnesota  herbaria. 


5.     I.  Harveyi   n.  sp. 

Trunk  deeply  2-  or  occasionally  3-lobed,  1.6-3  cm- in  diameter  : 
leaves  50-140,  short  (5-6cm.),  very  stout  and  fleshy,  2.5-3111111.  in  di- 
ameter, strongly  recurved,  with  an  abrupt  sharp  point,  purple-bronze 
in  color,  often  reddish  in  drying,  without  bast  or  stomata  :  ligula 
short,  obtuse  :  velum  ^-^3  indusiate,  sporangium  small,  unspotted: 
macrospores  526-648,  abnormally  810  ,a,  with  irregular,  parallel, 
thickish  crests  above,  and  honeycomb-reticulate  below,  more 
broken  in  large  spores  ;  microspores  35. 2-39.6,"  long,  and  22-30  fj. 
broad,  rough. 

I  have  hesitated  somewhat  whether  to  make  this  a  variety  of 
Tuckermani  on  its  spore  characters,  or  a  species  on  its  leaf  charac- 
ters, and  finally  concluded  that  it  is  just  as  much  entitled  to  specific 
rank  as  Tuckermani  itself,  which  in  spore  character  is  very  near 
riparia,  but  perfectly  distinct  in  leaf  character.  Although  in  all 
three  of  these  species  the  general  average  of  spores  have  an  indi- 
vidual appearance,  some  can  easily  be  found,  which,  if  placed  with 
selected  spores  of  the  other  species,  could  not  by  any  possibility  be 
separated.  The  group  seems  to  be  connected  with  lacustris  by 
riparia  in  spore  characters,  and  some  spores  of  Braunii  are  much 
like  some  spores  of  lacustris.  As  previously  remarked,  spore  char- 
acters are  a  very  unsafe  basis  for  characterization,  but  one  who  ex- 
amines many  sets  of  these  species  will  be  struck  with  the  gradation 
from  the  slender  spined  Boottii  to  the  beautifully  reticulate  Tuck- 
er n <iaui. 

The  leaves  of  Harveyi  are  relatively  the  stoutest  of  any  North 
American  species,  and  are  equalled  only  by  the  Peruvian  species, 
/  Lechleri  Mett.,  which  apparently  belongs  to  the  same  group. 

Pushaw  Pond,  Oldtown,  Maine,  F.  L.  Harvey;  pond  north  of 
Long  Pond,  Mt.  Desert,  Fernald  in  part. 

Type  in  the  herbarium  of  A.  A.  Eaton  ;  co-types  in  the  U.  S. 
National,  and  the  University  of  Minnesota  herbaria. 

6.     I.  RIPARIA    Engelm. 

Polygamous  :  a  medium  sized  species,  with  15-30  erect,  terete 
leaves,  io-3ocm.  long  :  sporangium  spotted  ;  velum  #-#  indusiate; 
macrospores  450-650^  in  diameter,  with  thin,  jagged,  irregular, 
mostly  short  crests,  sometimes  more  confluent  and  reticulate — al- 
most exactly  intermediate  in  sculpture  between  lacustris  and  Tuck- 
ermani. 

Reported  from  Maine,  but  all  specimens  so  labeled  which  have 


come  under  my  observation,  have  proven  to  be  Tuckermam  or  the 
variety  borealis,  notably  the  Mt.  Desert  locality  of  Redfield  and 
Rand's  Flora.  I  cannot  agree  with  Mr.  Dodge  in  his  East  King- 
ston, N.  H.,  nor  Newburyport,  Mass,  localities.  Also  reported 
from  Uxbridge,  Mass.,  and  Brattleboro,  Vermont.  It  is  extremely 
rare  in  New  England,  to  say  the  least,  and  the  specimens  from  the 
old  localities  should  be  verified. 

7.     I.  FOVEOLATA  A.  A.  Eaton. 

Trunk  2- or  rarely  3-lobed :  leaves  15-70,  very  stout,  5-iscm. 
long,  pink,  becoming  olive-green,  stomata  few  :  plants  apparently 
polygamous  :  sporangium  spotted ;  macrospores  380-560  //,  with 
lower  surface  full  of  little  holes,  made  by  the  close  reticulation  of 
the  thick  walls,  sculpture  of  upper  faces  more  open  ;  microspores 
22-35  (j.  long,  reticulated. 

Lamprey  (or  Pautuckaway)  river,  Epping,  and  Newmarket, 
N.  H.  Mr.  Dodge's  East  Kingston  locality  rests  upon  a  single 
plant. 

8.       I.   ECHINOSPORA     DurieU. 

Not  certainly  known  from  America,  our  varieties  all  having 
stomata,  while  the  species  does  not.  I  have  specimens  from  Brad- 
ley, Maine,  collected  by  Harvey,  which  I  have  examined  so  far  in 
vain  for  stomata.  They  may  prove  typical  in  this  respect,  but  dif- 
fer in  other  particulars. 

8a.     I.  ECHINOSPORA  BRAUNii  ( Durieu )  Engelm. 

Leaves  10-30,  usually  grass-green,  spreading,  5-23cm.  long ; 
stomata  few  if  immersed,  many  if  emersed :  sporangia  pale  spot- 
ted, velum  Yz  indusiate  ;  macrospores  350-550  n  in  diameter,  cov- 
ered with  short,  broad,  forked  spinules,  usually  mixed  with  longer 
slender  ones. 

As  commonly  understood,  this  variety  is  found  abundantly  in 
New  England  in  muddy  mill  ponds  and  river  banks,  also  in  sand  at 
times,  the  plant  then  usually  having  a  reddish  tinge,  and  the  leaves 
often  spiral,  especially  if  submersed.  Dodge  says  it  is  often  found 
where  the  water  is  very  brackish,  but  I  have  been  unable  to  verify 
this.  Though  subject  to  three  or  more  fest  of  tide  in  the  Merrimac 
at  Newburyport,  the  water  is  sweet  save  under  exceptional  condi- 
tions. I  am  unable  to  say  whether  the  same  conditions  prevail  in 
the  Maine  rivers  mentioned  by  him. 


—13— 

Our  species  of  Isoetes  appear  to  be  very  sensitive  to  salt  water, 
and  I  have  been  unable  to  find  any  under  its  direct  influence,  save 
a  few  plants  of  Engehnanni  in  a  mud  hole  near  a  dyke  next  the 
silt  marsh,  where  inundated  only  by  the  highest  tides. 

8b.     I.  ECHINOSPORA  ROBUSTA   Engelm. 

Stouter,  with  25-70  leaves,  bearing  more  stomata. 

So  far  as  herbarium  specimens  are  concerned,  this  differs  from 
llraunii  only  in  the  above  characters,  but  the  collector  who  has  a 
general  idea  of  Isoetes,  can  usually  determine  when  he  has  this  va- 
riety, by  its  appearance.  It  must  manifestly  differ  in  more  than  size, 
or  a  plant  might  be  Braunii  one  year  and  robusta  the  next.  Though 
it  is  impossible  to  separate  all  small  plants  with  certainty,  a  goodly 
percentage  can  be  so  separated  by  the  appearance  of  the  plants,  but 
it  is  difficult  to  satisfactorily  define  these  differences  in  a  descrip- 
tion. 

Lake  Champlain,  Pringle;  Epping,  N.  H.,  A.  A.  Eaton. 

8c.     I.  ECHINOSPORA  BooTTii  (A.  Br. )  Engelm. 

Leaves  io-i2.5cm  long,  stiffly  erect,  of  a  soft,  delicate  green  : 
spinules  of  spores  long  and  slender. 

Not  certainly  known  except  from  Boott's  original  localities  near 
Boston. 

8d.     I.  ECHINOSPORA  MURICATA  (Durieu)  Engelm. 

Leaves  10-15,  ver>"  slender,  the  submersed  ones  i5-4ocm.  long, 
ascending,  in  spirals,  stomata  few  :  spores  400-480  ft  with  slender 
spinules  often  mixed  with  short  or  elongated  crests. 

After  the  fall  of  the  water  in  summer,  the  long  leaves  disap- 
pear, and  are  succeeded  by  short,  bright  green  ones  5-7011.  long. 
In  this  state  the  plants  would  not  be  taken  for  the  same  unless  the 
transition  had  been  observed.  There  is  a  specimen  from  Boott's 
herbarium,  collected  in  this  stage,  in  the  National  Herbarium. 

Maine  :  Moosehead  Lake,  Harvey;  various  localities,  Fernald. 
New  Hampshire  :  Kingston,  East  Kingston,  Newton,  Epping,  New- 
market, A.  A.  Eaton.  Vermont:  Norwich  (fide  Dodge).  Massa- 
chusetts :  Woburn  creek  and  Abajona  river,  Boott.  Connecticut : 
Groton,  Graves. 

9.     I.  EATONI    Dodge. 

The  largest  New  England  species,  exceeded  in  size  by  none 
now  known  in  the  world,  and  equalled  only  by  Engelmanni  valida 


and  the  European  Maliaverniana.  Leaves  25-200,  as  much  as  6ocm. 
in  length  in  water,  about  io-i5cm.  when  growing  on  banks.  Known 
at  once  in  the  field  by  its  size,  but  especially  by  the  sporangium, 
which  is  light  brown  in  color  ( white  in  nearly  all  other  New  Eng- 
land species),  and  is  sparsely  rilled  with  very  small  spores.  Mic- 
rosporangia  are  rarely  found  in  this  species,  but  as  it  is  abundant  in 
several  localities,  it  certainly  must  bear  microspores,  as  it  never 
multiplies  by  offshoots.  It  may  be  possible,  but  not  probable,  that 
some  sporanges  bear  both  kinds  of  spores.  I  have  noted  such 
sporanges  in  Tuckermani  and  several  other  species,  but  the  micro- 
spores  are  usually  aborted  in  such  cases. 

Found  thus  far  only  in  the  waters  of  three  small  rivers — Powow 
at  East  Kingston  and  Kingston,  N.  H.,  and  Amesbury,  Mass., 
in  the  Lamprey  at  Epping  and  Newmarket,  A.  A.  Eaton;  in  Parker 
river  at  Georgetown,  Mass.,  Dodge. 

10.     I.  Gravesii  n.  sp. 

Plant  dioecious  or  polygamous,  rather  large  :  rootstock  bi- 
lobed  :  leaves  50-75,  12-15011.  long,  imm.  in  diameter  in  the  mid- 
dle, erect,  sharp  pointed,  dark  green,  with  abundant  stomata  and 
four  bast-bundles  :  velum  quite  narrow,  inner  sporanges  oval,  light 
cinnamon  in  color  from  the  abundance  of  vermiform,  translucent, 
light-colored  sclerenchym  cells  ;  macrospores  many,  small,  351-405  n 
in  diameter,  tetrahedro-globose,  the  upper  facies  flat,  densely  cov- 
ered with  short,  truncate,  mostly  single  columns  ;  microspores  not 
seen. 

Goshen,  Conn.,  Underwood,  1899;  gravelley  tidal  shore,  Sel- 
dens  Cove,  Lyme,  Conn.,  August  31,  1900,  Dr.  C.  B.  Graves. 
Specimens  were  sent  to  Dodge  from  this  locality  by  Graves  in  1895. 
It  was  at  first  referred  to  Eatoni,  but  was  finally  separated,  and  has 
since  lain  without  name.  It  has  the  aspect  of  sac  charata;  \s\\\.  its 
affinities  are  with  Eatoni,  with  which  it  agrees  in  being  polygamous, 
in  appearance  of  sporangium  and  shape  of  spores,  which  in  both 
have  the  appearance  of  being  abortive.  It  is  a  smaller  plant  with 
erect  leaves,  while  the  emersed  ones  of  Eatoni  are  not.  The  spore 
sculpture,  resembling  echinospora  rather  than  Eatoni,  safely  sepa- 
rates them.  Though  the  majority  of  the  columns  are  single,  each 
spore  usually  has  a  few  connected  into  vermiform  or  horseshoe- 
shaped  figures,  but  they  are  walls  rather  than  wrinkles,  and  in  this 
respect  resemble  Eatoni  but  remotely. 

The  type  is  in  my  private  herbarium  ;  co-types  are  deposited 


in  the  U.  S.  National  Herbarium,  in  the  herbaria  of  the  Missouri 
Botanical  Garden  and  the  University  of  Minnesota. 

ii.     I.  DODGEI   A.  A.  Eaton. 

Plants  medium  to  large  :  leaves  10-75,  the  submersed  spirally 
ascending,  2o-45cm.  long,  the  emersed  io-2ocm.  long,  often,  espe- 
cially when  the  plants  are  not  crowded,  tortuous  and  interlaced, 
2-3mm.  broad  :  velum  one-fifth  to  one-fourth  indusiate  ;  sporangium 
spotted  ;  macrospores  500-675  /*,  averaging  560  p,  with  mostly  scat- 
tered groups  of  irregular,  spinulose-rosulate  crests  ;  microspores 
ashy,  22-40  /u,  wrinkled. 

Abundant  on  the  banks  of  Powow  river  where  overflowed 
for  the  purpose  of  forming  a  pond  during  the  greater  part  of  the 
winter,  at  Kingston,  N.  H.,  the  only  known  New  England  locality. 

12.     I.  ENGELMANNI    A.  Br. 

Leaves  15-100,  bright  green,  usually  erect  except  when  grow- 
ing out  of  water  in  bare  places,  stomata  abundant :  sporangium  un- 
spotted, velum  narrow  ;  macrospores  chalk-white,  350-550 /</,  honey- 
comb-reticulated ;  microspores  24-29  fi,  smooth. 

Very  common  in  clayey  soil  in  ponds  and  ditches,  rarely  in 
mud  or  sand.  For  the  greater  part  of  the  season  it  is  likely  to  be 
obscured  by  a  tangle  of  swamp  grasses  and  other  vegetation. 

The  spores  are  peculiar,  those  of  no  other  American  species 
approaching  them  in  sculpture.  Those  of  Tuckermani,  riparia, 
hieroglyphica,  and  Harveyi  are,  it  is  true,  more  or  less  honey- 
combed for  a  part  of  the  surface,  but  the  pattern  is  always  incom- 
plete. The  nearest  approach  to  it  is  Japonica,  in  which  the  spores 
are  marked  almost  identically,  but  the  crests  are  shorter  and  less 
delicate,  and  the  reticulations  larger,  as  well  as  the  spores. 

In  Duriaei  of  Europe  the  pattern  is  also  the  same,  but  the  walls 
are  ridges  rather  than  laminae,  and  the  spores  double  the  diameter. 
In  Azorica  the  spores  are  similar  in  size,  but  the  reticulations  are 
less  regular,  the  walls  low,  and  there  is  little  of  the  honeycomb. 
The  only  othej  species  with  spores  of  this  pattern,  is  Schweinfurthii 
from  equatorial  Africa.  My  specimens  of  this  species  are  immature, 
but  one  spore  would  pass  for  Azorica,  while  another  is  covered 
with  short,  blunt,  crowded  tubercles  or  ridges.  The  latter  was 
taken  from  the  roots,  and  may  be  intrusive. 

In  specimens  from  farther  south  the  spores  have  a  tendency  to 
lose  their  characteristic  form,  and  the  species  breaks  up  into  vari- 


eties.     I  have  seen  no  typical  Engehnanni  from  south  of  Pennsyl- 
vania. 

Maine  :  Fernald,  according  to  Dodge,  though  there  are  no 
specimens  in  Fernald's  collection,  which  he  has  kindly  sent  to  me 
for  examination.  New  Hampshire  :  in  nearly  every  mill  pond  or 
ditch  with  clay  bottom  in  Rockingham  county.  Massachusetts : 
common  in  the  eastern  part.  Rhode  Island  :  Newport,  Farloic,  fide 
Dodge.  Connecticut :  Meriden,  Hall;  Waterford  and  Groton, 
Graves. 

i2a.    I.  ENGELMANNI  GRACILIS   Engelm. 

An  attenuate  form  of  the  type,  and  often  growing  with  it ;  ap- 
parently caused  more  by  environment  than  any  inherent  qualities. 

Seabrook,  N.  H.,  June  27,  1900. 


THE  SYSTEJ1  OF  FERNS  PROPOSED  IN  DIE 
NATUERLICHEN  PFLANZENFAHILIEN. 

Bv  LUCIEN  M.  UNDERWOOD. 


To  those  familiar  only  with  the  system  of  Fern  Genera 
that  has  been  followed  in  Synopsis  Fi/icum,  and  in  its  main 
features  has  been  accepted  in  its  application  to  the  ferns  of 
North  America  by  most  who  have  made  our  ferns  a  study,  the  sys- 
tem proposed  in  Engler  &  Prantl's  Die  Natiterlichen  Pflanzenfa- 
milien  will  present  many  strange  features  ;  yet,  on  the  whole,  it  is 
to  be  regarded  as  a  very  conservative  arrangement.  Nevertheless, 
while  the  true  ferns  (Polypodiaceae)  of  Synopsis  Filicum  number 
only  47  genera,  the  same  group  in  the  present  arrangement  is  dis- 
tributed among  109  ;  and  were  the  system  as  uniformly  consistent 
as  it  is  in  places,  the  number  would  quite  readily  be  increased  to 
three  times  that  of  Synopsis  Filicum.  Mettenius,  Kuhn,  and  Prantl, 
the  three  great  German  fern  taxonomists,  have  all  passed  away, 
and  it  was  left  to  a  novice  among  ferns — but  none  the  less  a  trained 
botanist — to  bring  the  genera  together ;  it  is  to  be  judged  then  in 
this  light,  and  not  as  the  result  of  a  long  continued  personal  study. 
While  it  is  a  great  improvement  on  the  system  that  has  too  long 
been  in  vogue  as  the  expression  of  a  part  of  the  obsolete  English 
school  of  fern  taxonomists,  it  is  still  lacking  in  many  of  the  charac- 
teristics and  consistencies  that  a  genuine  system  must  possess. 


—17— 

XYhile  some  of  the  excesses  of  the  systems  of  Fee  and  Moore  are 
not  accepted,  many  of  the  really  scientific  aspects  of  the  systems  of 
Presl  and  John  Smith  have  not  been  incorporated,  and,  as  a  whole, 
it  lacks  most  what  a  master  would  have  put  into  it — homogeneity 
and  consistency  of  treatment.  For  a  system  that  is  supposed  to 
proceed  from  low  to  high  in  an  evolutionary-  way,  the  order  of  ar- 
rangement of  the  larger  groups  is  surely  peculiar,  as  may  be  seen 
by  the  following : 

Family  HYMENOPHYLLACEAE 
Family  CYATHEACEAE 
Family  POLYPODIACEAE 

1.  IVoodsieae 

2.  Aspidieae 

3.  Oleandreae 

4.  Davallieae 

5.  Asplenieae 

6.  Pterideae 

7.  Viitarieae 

8.  Polypodieae 

9.  Acrosticheae 
Family  PARKERIACEAE 
Family  MATONIACEAE 
Family  GLEICHENIACEAE 
Family  SCHIZAEACEAE 
Family  OSMUNDACEAE 

The  family  OPHIOGLOSSACEAE  is  removed  to  a  separate  order, 
as  is  also  the  family  MARATTIACEAE. 

\Yhile  there  is  long  likely  to  be  two  schools  of  belief  regarding 
the  relative  position  of  the  eusporangiate  and  leptosporangiate 
forms,  it  is  quite  evident  that  were  the  groups  of  the  Polypodiaceae 
inverted,  they  would  come  nearer  representing  an  ascending  series, 
and  some  of  the  later  families  are  surely  simpler.  As  to  these 
major  groups  themselves,  there  is  little  fault  to  be  found  ;  possibly 
Platycerium  might  be  held  by  some  to  a  more  distinctive  rank  than 
simply  a  mere  member  of  the  tribe  Acrosticheae,  but  in  the  main 
the  separation  into  families  is  logical  and  scientific.  With  the  ex- 
ception of  Parkeriaceae,  the  names  are  well  chosen  ;  this  one  is  un- 
fortunate as  based  on  one  of  the  many  synonyms  of  Ceratopteris, 
while  the  name  Ceratopteridaceae  which  is  possibly  less  euphonious 
has  the  double  merit  of  being  distinctive,  and  (in  one  of  its  forms 
at  least)  more  ancient.  Some  of  the  commendable  features  of  the 
work,  so  far  as  they  pertain  to  our  American  ferns,  may  be  noted 
in  brief  form  : 


1.  The  separation  of  Dennstaedtia  from  Dicksonia,  and  the 
breaking  up  of  the  latter  genus  into  three.     Our  own  species  of 
Dennstaedtin,  too  long  held  as  a  Dicksonia,  is  separated  from  that 
genus  not  only  generically  but  family-wise. 

2.  The  separation  of  our  two  species  hitherto  combined  under 
Onoclea  into  two  genera, — certainly  as  distinct  genera  as  were  ever 
created  ! 

3.  The  breaking  up  of  the  aspidioid  forms  into  several  genera. 
While  this  has  not  always  been  as  thorough  as  might  have  been 
wished,  it  is  a  good  beginning.     Of  course  the  selection  of  generic 
names  was  bound  to  be  unfortunate,  since  the  work  is  presumably 
based  on  the  absurd  and  illogical  fifty  year  system.     Personally  I 
am  loth  to  see  such  genera  as  Phegopteris,  Meniscium,  Goniopteris, 
and  the  like,  swallowed  up  with  Dryopteris,  but  I  can  admit  the 
presence  of  connecting  forms,  which  is  no  more  than  we  as  evolu- 
tionists must  expect  in  genera  as  well  as  species. 

4.  Altho  American  only  in  cultivation,  the  breaking  up  of  Da- 
vallia  into  many  genera  should  be  commended.     The  transference 
of  Nephrolepis  to  the  Davallieae  is  also  logical. 

5.  The  separation  of  Asplenium  into  several  genera.     Here 
again  the  division  could  consistently  be  made  still  greater. 

6.  The  breaking  up  of  the  composite  genus  Gymnogramma 
as  hitherto  understood. 

7.  The  removal  of  Notholaena  (wrongly  written  Nothochlaena~] 
to  a  position  intermediate  between  Pellaea  and  Cheilanthes. 

8.  The  removal  of  Pellaea  Stelleri  to  Cryptogromma,  follow- 
ing Prantl's  lead. 

9.  The  breaking  up  of  Pteris  into  several  genera,  of  which 
our  Pteris  aquilina  becomes  Pteridium. 

10.  The  division  of  Taenitis  into  several  genera. 

11.  The  breaking  up  of  Acrostichum  as  comprised  under  the 
Hookerian  (Kew)  system  into  several  genera,  of  which  our  species 
properly  remain  under  Acrostichum  itself. 

These  are  some  of  the  many  favorable  points  in  the  Berlin  sys- 
tem.    Among  the  questionable  points  we  may  note  the  following  : 

1.  The  retention  of  Camptosorus  and  Scolopendriuni  in  one 
genus — as  has  always  been  maintained  in  the  Kew  system. 

2.  The  union  of  Lomaria  and  Blechnum.     Despite  the  resem- 
blance, there  seems  to  us  a  real.difference. 

3.  The  joining  of  certain  species  of  Notholaena  like  nivea  and 
tenera  to  Pellaea.     Possibly  a  separation  of  each  genus  into  two 


—19— 

would  be  more  natural,  but  this  deserves  a  study  of  a  wide  array  of 
forms. 

4.  The  unfortunate  and  illogical  treatment  of  Polypodium.  A 
few  of  the  more  striking  groups  like  Dipteris,  Drynaria,  and  Ar- 
thropteris  have  been  separated,  but  this  is  only  a  beginning.  Of 
our  American  species,  P.  polypodioides,  the  least  divergent  of  all, 
has  been  placed  in  the  genus  Lepicystis,  while  the  others  are  all  re- 
tained in  Polypodium.  And  the  same  characters  that  elsewhere 
have  been  regarded  as  of  importance,  namely,  vegetative  charac- 
ters, have  been  utterly  disregarded.  Surely  our  Polypodium  vul- 
gare,  aureum,  phyllitidis,  and  Swartzii  represent  four  distinct  gen- 
era if  Diplazium  is  to  be  separated  from  Asplenium,  Polystichum 
from  Dryopteris,  and  Camptosorus  from  Scolopendrium,  or  either 
from  Asplenium,  where  Linnaeus  placed  them. 

Venation,  belogning  to  a  fundamental  portion  of  the  growth  of 
the  sporophyte,  must  for  the  future  be  regarded  as  of  first  import- 
ance in  the  taxonomy  of  ferns,  and  when  the  system  of  the  future  is 
finally  attained,  we  believe  it  will  be  found  that  the  systems  of  Presl 
and  John  Smith  will  approximate  very  closely  to  that  ideal.  This 
will  involve  a  greater  regard  for  the  fibre-vascular  system  as  repre- 
sented in  venation,  and  growth  characters  based  on  a  study  of  root- 
stock  and  caudex,  than  we  have  presented  in  the  system  under  re- 
view. 

Columbia  University,  June,  1900. 


EXPERIMENTS   IN   HYBRIDIZING  FERNS. 

BY  MARGARET  SLOSSON. 


Various  arguments  have  been  advanced  both  for  and  against 
the  probable  occurrence  of  hybridity  among  ferns  in  nature,  but  I 
shall  not  here  enter  into  the  "pros  and  cons."  There  is  now  no 
doubt  that  fern  hybridization  is  possible  ;  it  has  been  brought  about 
artificially.  To  give  the  most  convincing  instance,  Mr.  E.  J.  Lowe 
has  succeeded  in  crossing  Ceterach  officinarum  and  Scolopendrium 
vulgare,  two  ferns  belonging  not  only  to  different  species,  but  to 
different  genera. 

The  next  step  is  to  ascertain  whether  fern-hybridization  occurs 
in  nature  ;  if  so,  to  what  extent,  and  which  are  the  hybrid  ferns. 


Obviously,  the  only  infallible  way  to  discover  this,  is  to  cross  the 
supposed  parents  of  suspected  hybrids,  and  compare  with  the  latter 
the  resultant  plants.  All  other  evidence  regarding  suspected  hy- 
brids, such  as  abortion  or  eccentricity  of  the  fronds,  sterility  of  the 
spores,  rarity  of  the  plants,  presence  of  the  supposed  parents,  while 
pointing  to  the  almost  irresistible  conclusion  that  the  fern  is  a  hy- 
brid, is  not  positive  proof.  I  have  found  no  record  of  any  attempt 
of  the  kind.  American  pteridologists  appear  to  have  left  the  hy- 
bridizing of  ferns  to  pteridologists  abroad,  who  have  confined  their 
attention  to  producing  either  entirely  new  forms,  or  new  varieties 
of  the  crested  and  laciniate  ferns  so  dear  to  our  English  cousins. 

I  am  indebted  to  Mr.  George  E.  Davenport's  paper  on  "  Hy- 
bridity  in  Ferns,"  read  before  the  Boston  meeting  of  the  Chapter, 
not  only  for  calling  my  attention  to  the  subject,  but  also  for  sug- 
gesting the  experiments  I  have  tried.  In  order  to  make  clear  a  de- 
scription of  these  experiments,  it  will  be  necessary  for  me  to  repeat 
part  of  what  he  has  already  said. 

There  are  several  methods  by  which  the  crossing  of  the  sup- 
posed parents  of  a  suspected  fern-hybrid  may  be  attempted.  As- 
suming two  species  to  be  capable  of  blending,  and  their  prothalli 
monoecious,  I  will  state  briefly  the  possibilities  of  failure  in  each 
method. 

The  direct  crossing  of  ferns  by  the  artificial  transference  of  an- 
therozoids  from  one  prothallus  to  the  archegonium  of  another  is 
very  difficult.  I  believe  no  one  has  succeeded  in  accomplishing  it. 
I  have  tried  and  failed.  Owing  to  the  infinitesimal  size  of  the  an- 
therozoids,  it  must  be  done  under  a  microscope  and  some  instru- 
ment must  be  devised  that  will  lift  the  antherozoids — not  quiescent 
and  adhesive  like  pollen,  but  moving  rapidly — and  deposit  them 
without  injury  upon  the  desired  archegonium. 

Other,  less  direct,  methods  have  proved  more  successful.  The 
simplest  of  these  is  to  sow  a  mixture  of  the  spores  of  the  two  ferns, 
and  trust  to  the  proximity  of  the  resultant  prothalli  to  bring  about 
cross-fertilization.  The  disadvantages  of  this  method  are,  that  it  is 
impossible  to  tell  whether  the  germinating  spores  come  from  both 
species  (which  introduces  an  element  of  doubt  into  the  final  result), 
that  all  or  nearly  all  of  one  species  may  fail  to  germinate,  and  that, 
in  order  not  to  lose  a  possible  hybrid,  it  is  necessary  to  raise  every 
one  of  the  young  plants,  which  means  a  great  waste  of  time  and 
trouble.  The  objection  may  also  be  urged  that  even  should  both 
species  germinate,  one  might  germinate  later  or  develop  faster  than 


the  other,  and  thus  none  of  the  prothalli  of  both  ripen  for  fertiliza- 
tion at  the  same  time,  but  I  think  this  would  rarely  happen.  I  have 
found  with  every  species  I  have  raised  from  spores,  that  while  many 
of  the  prothalli  develop  plants  at  the  same  time,  usually  on  the 
same  day,  others  remain  apparently  latent  for  long  periods,  some- 
times for  months. 

A  third  method,  successfully  used  by  Professor  Bower  in  cross- 
ing Polvpodiuui  anreum  and  a  variety  of  Polypodium  vulgare,  is 
that  of  sowing  the  spores  of  each  species  separately,  afterward 
planting  a  prothallus  of  one  species  in  close  contact  with  a  prothal- 
lus  of  the  other.  This  method  offers  more  certainty  of  success,  but 
allows  each  species  to  retain  both  kinds  of  organs,  which  gives  ex- 
cellent opportunity  for  self-fertilization  as  well  as  for  cross-fertiliza- 
tion. 

A  fourth  method,  pursued  by  Mr.  Lowe,  is  that  of  sowing  the 
spores  separately,  afterward  cutting  the  resultant  prothalli  into  quar- 
ters and  planting  a  quarter  of  a  prothallus  of  one  species  overlap- 
ping a  quarter  of  a  prothallus  of  the  other.  In  order  that  this 
method  may  succeed,  it  is  of  course  necessary  that  one  of  these 
quarters  shall  contain  archegonia  and  the  other  antheridia.  Cutting 
the  prothalli  in  this  way  would  probably  as  a  rule  isolate  each  kind 
of  organ.  The  danger  of  this  method  lies  in  the  fact  that  the  pro- 
thallus is  so  small  that  by  the  slightest  slip  of  the  knife  one  might 
accidentally  leave  a  bit  of  tissue  containing  archegonia  on  an  anther- 
idial  section,  or  vice  versa. 

A  fifth  method,  suggested  by  Mr.  Davenport,  and,  so  far  as  I 
know,  not  tried  by  anyone  heretofore,  is  that  of  sowing  the  spores 
separately,  cutting  the  resultant  prothalli  in  two  between  arche- 
gonia and  antheridia,  then  planting  an  archegonial  section  of  one 
species  in  contact  with  an  antheridial  section  of  the  other,  so  that 
archegonia  and  antheridia  coalesce.  This  is  the  methodl  have  used, 
with  the  slight  modification  of  cutting  the  prothallus  into  three  sec- 
tions instead  of  two  ;  cutting  once  transversely  close  to  the  sinus 
and  through  the  cushion  of  tissue  on  which  the  archegonia  in  most 
species  are  borne  ;  cutting  again  transversely  near  the  base,  through 
the  root-hairs  among  which  the  antheridia  are  scattered,  then  dis- 
carding the  middle  section  and  planting  the  upper  (archegonial) 
section  of  one  species  upright  against  the  lower  (antheridial)  sec- 
tion of  the  other,  with  the  lower  surfaces  of  both  sections  pressed 
close  together.  Cutting  in  this  way  prevents,  I  think,  all  danger  of 
leaving  both  kinds  of  organs  on  the  same  section.  There  is  some 


danger  that  the  prothalli  may  be  cut  too  late,  after  fertilization  has 
taken  place,  in  which  case  plants  of  pure  species  would  develop. 
Since  some  prothalli  when  fully  grown  are  barely  one-eighth  the  size 
of  others,  while  many  grow  at  varying  rates  of  speed,  it  will  be 
found  impossible  when  cutting  to  distinguish  between  the  small  un- 
developed prothalli  and  those  which  though  small  are  already  fer- 
tilized, except  perhaps  by  the  use  of  a  powerful  microscope.  How- 
ever, the  newly  cut  archegonial  sections,  before  being  brought  in  con- 
tact with  antheridial  sections  of  the  other  species,  could  be  isolated 
long  enough  to  allow  all  fertilized  sections  to  develop  plants  and  be 
thrown  away.  There  is  also  danger  that  plants  may  develop  at  any- 
time from  asexual  growth,  such  as  Dr.  Farlow  discovered  in  Pteris 
cretica.  But  Dr.  Farlow  has  pointed  out  that  in  plants  arising  in  this 
way,  the  first  frond  springs  directly  from  the  prothallus,  so  that  it  is 
impossible  to  tell  where  one  begins  and  the  other  ends,  and  the  first 
root  develops  afterward  from  the  base  of  this  frond  ;  while  in  plants 
arising  from  fertilized  archegonia,  according  to  Sachs,  both  frond 
and  root  are  separated  from  the  prothallus  by  a  slight  connectile 
known  as  the  "foot,"  and  the  first  root  develops  from  the  base  of 
this  foot  some  time  before  the  appearance  of  the  first  frond  from  its 
apex.  Thus  plants  arising  from  asexual  growth,  as  well  as  from 
self-fertilization  previous  to  the  cutting,  could  be  detected  and 
weeded  out. 

One  would  naturally  suppose  that  all  other  plants  arising  from 
these  sections  must  necessarily  be  the  result  of  cross-fertilization. 
Unfortunately,  there  is  doubt  of  that.  The  prothallial  sections  usu- 
ally branch  soon  after  being  cut.  They  nearly  always  do  so  sooner 
or  later.  The  antheridial  sections  often  die  down  for  several  weeks, 
and  then  send  up  whole  clusters  of  prothalli.  Or  new  segments 
may  appear  from  any  part  of  the  margin  of  the  old.  In  archegonial 
sections  a  new  segment  frequently  fills  up  the  old  sinus.  Sometimes 
the  effect  is  that  of  an  old  prothallus  with  young  prothalli  starting 
from  the  edge  ;  sometimes  of  a  cluster  of  overlapping  prothalli 
continuous  with  one  another  at  the  base.  Each  new  segment  is 
shaped  like  a  complete  prothallus,  sinus  and  all,  which  naturally 
suggests  the  question,  may  not  these  new  segments  replace  the 
organs  that  have  been  cut  away  ?  I  have  not  yet  discovered  whether 
this  occurs  or  not.  Mr.  Lowe's  experience  in  quartering  prothalli 
would  tend  to  show  that  it  does  not.  He  speaks  of  the  branching 
of  the  quarters,  and  states  that  while  keeping  the  lower  quar- 
ters in  one  dish  and  the  upper  in  another,  no  plants,  with  one  ex- 


—23— 

ception,  appeared.  This  exceptional  plant  may  have  been  the  result 
of  asexual  growth.  Mr.  Lowe  attributes  it  to  an  insect  carrying  an- 
therozoids  from  a  distance,  but  there  is  not  time  to  discuss  this  pos- 
sibility. If  the  sections  do  replace  the  missing  organs,  this  fifth 
method,  as  well  as  all  the  others  except  the  first  described,  can  be 
used  for  testing  the  hybridity  of  a  fern  only  in  cases  where  the 
alleged  hybrid  is  so  distinct  from  both  of  its  supposed  parents  that 
it  could  not  possibly  arise  from  the  self-fertilization  of  either  one. 

Of  such  a  nature  are  Asplenium  ebenoides  and  Aspidium  cris- 
tatum  x  marginale.  I  suppose  no  one  will  question  for  a  moment 
that  Asplenium  ebenoides  cannot  be  considered  a  variety  of  either 
Asplenium  ebeneum  or  Camptosorus  rhizophyllus,  but  must  either 
be  a  cross  between  the  two  or  a  distinct  species.  Now  a  distinct 
species  certainly  could  not  arise  from  a  prothallus  of  another  species, 
either  from  self-fertilization  or  asexual  growth.  Therefore  if  a  plant 
of  ebenoides  should  arise  from  a  prothallus  of  ebeneum  or  Campto- 
sorus, ebenoides  must  be  a  hybrid.  Moreover,  the  characters  of 
ebeneum  and  Camptosorus  are  such  that  a  hybrid  between  the  two, 
even  should  it  prove  unlike  ebenoides,  must  be  plainly  distinguish- 
able from  plants  of  pure  species  of  either  fern.  And  the  same  is 
true  of  Aspidium  cristatum  x  marginale  and  its  supposed  parents, 
marginale  and  cristatum.  This  fifth  method  offers,  then,  a  good 
way  of  obtaining  positive  proof  of  the  hybridity  or  non-hybridity  of 
Asplenium  ebenoides  and  Aspidium  cristatum  x  marginale.  It  of- 
ers,  also,  more  certainty  of  success  than  any  of  the  other  methods, 
whether  the  missing  organs  are  reproduced  or  not,  since  before  that 
could  occur,  cross-fertilization  would  probably  take  place. 

In  December,  1898,  with  a  view  to  testing  the  alleged  hybridity 
of  Aspidium  cristatum  x  marginale,  I  sowed  in  separate  flower  pots 
spores  of  marginale  and  of  cristatum,  first  sterilizing  the  earth  to 
destroy  all  alien  spores.  The  following  July,  the  resultant  prothalli 
had  grown  sufficiently  large  to  be  divided.  I  then  cut  and 
planted  archegonial  sections  of  cristatum  in  another  pot,  and  a 
week  or  so  later,  about  the  5th  of  August,  pressed  close  against 
them,  antheridial  sections  of  marginale.  The  sections  grew  and 
branched.  In  the  conglomeration  resulting,  it  was  soon  impos- 
sible to  tell  which  prothalli  belonged  to  marginale  and  which  to 
cristatum.  On  the  iyth  of  October,  a  plant  developed,  followed 
before  December  i3th  by  several  others.  Three  of  these  survived. 
One  of  the  three  grew  from  a  segment  of  a  prothallus  cluster.  Pivy 
months  afterward,  in  May,  1900,  seven  more  plants  developed  from 


—24— 

what  appeared  to  be  other  segments  of  the  same  cluster,  each  plant 
from  a  segment.  The  second  frond  of  two  of  these  seven  plants  is 
plainly  dichotomous,  a  frond  of  another  is  slightly  eccentric,  the 
other  fronds  are  normal.  One  of  the  three  earlier  plants  has  from 
the  first  shown  a  striking  tendency  to  produce  abortive  fronds,  and, 
so  far  as  one  can  tell  from  plants  barely  an  inch  and  a  half  high,  ap- 
pears truly  intermediate  between  young  typical  plants  of  marginale 
and  of  cristatum,  as  does  a  second  of  the  plants  developed  last 
fall.*  This  plant  has  produced  one  abortive  and  one  markedly  ec- 
csntric  frond.  The  fronds  of  the  third  of  last  fall's  plants  are  all 
normal  and  the  character  of  the  plant  is  like  that  of  pure  cristatum. 
In  May,  about  the  time  that  the  seven  later  plants  appeared,  eight 
more  developed  from  other  prothalli  in  the  same  pot.  All  have 
normal  fronds.  These  later  plants  cannot  be  the  result  of  self- 
fertilization  before  the  cutting;  the  time  that  has  elapsed  is  too 
great,  and  the  segments  from  which  many  or  all  have  sprung  were 
not  then  in  existence.  I  have  not  cared  to  disturb  them  in  order  to 
look  for  evidence  of  asexual  growth,  but  it  is  hardly  likely  that  all 
fifteen  have  arisen  in  that  way.  What  they  will  prove  to  be,  may 
throw  some  light  on  the  question  of  whether  or  no  the  missing 
organs  are  reproduced. 

In  order  to  obtain  a  reciprocal  cross,  I  cut,  in  October,  1899, 
archegonial  sections  of  marginale  and  antheridial  sections  of  cris- 
tatum. Because  the  time  was  so  long  since  the  spores  had  germi- 
nated, I  feared  that  some  of  the  archegonia  might  have  become  fer- 
tilized, and  so  isolated  both  sections  for  a  time  in  separate  pots. 
Four  plants  developed  within  a  month  and  were  thrown  away. 
Since  no  more  plants  appeared,  I  planted,  in  December,  the  two 
kinds  of  sections  together.  Both  kinds  had  branched  freely.  In 
March,  two  segments  of  one  section  developed  what  appeared  to  be 
a  broad,  much  thickened,  archegonial  cushion  of  tissue,  covered 
with  a  mucilaginous  liquid.  This  soon  dried,  and  the  cushions 
swelled  outward  into  points.  The  point  in  one  cushion  devel- 
oped into  a  horizontal  cylindrical  process  that  is  still  lengthening, 
and  has  flattened  at  the  end  into  a  fan-shaped  lobate  growth.  A 


*  These  two  plants  have  since  shown  marked  evidence  of  hybridity.  Their 
rootstocks  are  caudiciform,  like  rootstocks  of  cristatum  y.  marginale  Davenp. 
The  fronds  of  one  are  a  mean  between  those  of  its  parents.  Of  four  fronds  at 
present  011  the  other,  one  has  spinulose  margins,  like  young  cristatum.  one 
large  blunt  lobes,  like  young  marginale,  while  the  remaining  two  are  interme- 
diate in  form. 

December  6,  1900. 


—25- 
month  ago,  in  May,  1900,  four  plants  appeared  from  the  other  pro- 
thallial  sections  in  this  pot.  The  fronds  are  as  yet  small,  all  nor- 
mal, and  it  is,  of  course,  impossible  to  say  what  they  will  be.  Two 
other  attempts  at  crossing  cristatum  and  marginale  have  failed  to 
produce  plants. 

I  have  been  unfortunate  in  my  attempts  to  cross  Aspleniwn 
ebeneuin  and  Caniptosorus  rhizophyllns.  Prothalli  resulting  from 
spores  repeatedly  sown  in  sterilized  earth  have  repeatedly  died. 
But  I  have  now  several  archegonial  sections  of  Camptosorus,  and 
antheridial  sections  of  ebeneum,  which  were  cut  and  planted  to- 
gether last  winter.  One  of  the  archegonial  sections  has  produced 
a  peculiar  growth  which  seems  to  be  a  proliferous  bud  of  some 
kind  ;  the^  other  sections  have  branched  and  are  doing  well. 

Whether  plants  will  arise  from  these  and  prove  a  hybrid,  and 
that  hybrid  ebenoides,  time  alone  will  show. 

Andover,  Mass.,  June  27,  1900. 


ATHYRIUri    AS    A    GENUS. 

Bv  B.   D.  GILBERT. 


During  the  last  year  I  have  been  engaged  in  a  special  study  of 
the  group  of  ferns  classified  throughout  the  last  half  century  or 
more  as  Asplciiini>i  ftli.r-foemina.  In  the  course  of  this  study  it 
seemed  natural  that  I  should  take  up  also  the  question  of  the  genus 
of  these  ferns  ;  and  having  done  so,  I  propose  to.  lay  the  results  of 
that  study  before  the  Fern  Chapter,  as  briefly  as  the  subject  will 
permit. 

Although  all  writers  on  ferns  are  obliged  to  recognize  Athyrium 
by  the  character  of  its  sori,  the  tendency  has  been,  since  the  publi- 
cation of  Hooker's  Species  Filicum,  to  make  it  a  section  or  sub- 
genus  of  Aspleniuin.  Previous  to  Hooker's  time.  Roth's  genus 
Athyrium  had  been  quite  generally  accepted  by  German  and  even 
by  English  botanists ;  and  since  Hooker's  time,  Thomas  Moore, 
the  most  careful  monographer  of  the  English  ferns,  has  retained 
Athyrium  for  filix-foemi na  and  its  varieties.  In  our  own  country, 
since  the  time  of  Pursh,  who  followed  the  Linnaean  nomenclature, 
and  placed  these  ferns  under  Polypodiutn,  and  Dr.  Jacob  Bigelow, 
who  adopted  the  nomenclature  of  Swartz  and  regarded  them  as 


—26— 

forms  of  Aspidium,  American  botanists  have  accepted  Hooker's 
\-ie\vs.  Because  the  sori  are  long  and  the  indusia  are  attached  by 
one  side  to  the  veins  and  open  on  the  other  side,  all  of  which  fea- 
tures are  characteristic  of  Asplenium,  it  has  been  claimed  thaty?/T.r- 
foemina  and  other  species  allied  with  it  belong  to  that  genus.  But 
there  is  another  character  which  essentially  modifies  this  judgment 
and  compels  recognition.  That  is  the  curvature  of  the  sorus  and 
indusium,  by  which  the  upper  end  of  the  sorus  is  curved  across  the 
veinlet  and  back  upon  itself,  often  so  strongly  as  to  resemble  the 
sorus  and  indusium  of  a  typical  Lastrea,  or  free-veined  form  of 
\ephrodium,  distinguished  by  its  kidney-shaped  indusium.  It  was 
this  roundish  form  which  induced  Linnaeus  to  rank  it  as  a  I'olypo- 
dium,  and  Swartz  to  place  it  under  Aspidium.  These  three  genera 
therefore  have  laid  claim  to  it,  with  each  of  which  it  agrees  in  a 
modified  degree,  but  with  no  one  of  which  it  agrees  entirely.  The 
genus  Athyrium,  however,  as  constituted  by  Roth,  comprises  all 
these  features,  and  completely  satisfies  all  the  generic  conditions 
dependent  upon  its  fructification.  What  those  conditions  are  will 
best  be  discovered  by  taking  Roth's  own  description  of  the  genus, 
which  I  have  translated  from  the  Latin  : 

ATHYRIUM. 

"  Capsules  distributed  in  ovate  sori  underneath  the  disc  of  the 
frond,  surrounded  with  an  articulate  ring.  The  involucre  springs 
laterally  from  the  venule,  lying  loosely  in  the  form  of  a  scale,  with 
laciniate-fimbriate  margin,  at  length  elevated  inwardly,  pressed 
back  and  semi-lunar. 

"  Observation. — The  essential  character  of  Polystichum  consists 
in  the  involucre  being  either  umbilicate  or  peltate,  or  reniform,  but 
on  every  side  nearly  free.  In  the  first  case,  at  the  time  of  maturity, 
the  involucre  is  drawn  together  centrally  to  its  own  fixed  point,  and 
very  often  acquires  the  shape  of  a  funnel ;  but  in  the  latter  case  it  is 
drawn  back  sublaterally  to  a  fixed  point  and  changes  the  sub-peltate 
shape  into  a  reniform  shape. 

"  But  among  the  Linnaean  Polypods  there  is  observed  no  other 
of  which  the  involucre  has  an  ovate-oblong  shape,  and  as  in  Asplen- 
iniu,  springing  laterally  from  the  venule,  draws  itself  out  following 
the  length  of  the  same.  It  lies  more  loosely  in  a  heap  before  the 
maturity  of  the  capsule,  when  from  the  other  side  opposite  and 
looking  backward  it  rises  a  little  above  the  costa  of  the  frond  or  of 
the  lacinia.  Toward  maturity,  it  is  raised  against  the  costa  by  the 


inward  increase  of  the  capsules  bursting  forth,  by  which  means  it 
assumes  a  semi-lunar  shape.  Influenced  by  these  reasons,  accord- 
ing to  the  method  of  founding  the  genera  of  ferns  upon  the  invo- 
lucre, I  have  come  to  the  conclusion  that  plants  of  this  nature  should 
be  removed  from  the  rest  of  the  Linnaean  Polypods,  on  account  of 
the  situation,  shape  and  condition  of  the  involucre.  Nor  can  they 
be  included  in  Asplenium,  on  account  of  the  plainly  distinct  condi- 
tion of  the  involucre,  although  they  approach  nearer  to  this  genus. 
Therefore  it  seems  necessary  that  I  should  establish  a  genus  of  their 
own,  to  which  I  have  given  the  name  Athyrium.'" 

Now  it  will  be  seen  from  this  description  that  Roth  distinctly 
differentiates  Athyrium  from  Asplenium.  He  places  only  seven 
species  under  the  genus,  of  which  A,  fontanum  and  A.  Halleri,  now 
recognized  as  forms  of  one  and  the  same  species,  are  placed  first. 
In  this  he  follows  the  universal  custom  of  placing  at  the  head  of  the 
list  the  smallest  and  simplest  forms,  not  because  they  are  the  most 
typical  species  of  the  genus,  but  because  they  are  the  smallest. 
The  next  five  species  are  all  now  recognized  as  varieties  of  A.  filix- 
foemina,  and  these  are  the  ones  which  conform  most  clearly  to  his 
description  of  the  genus. 

In  commenting  upon  this  genus,  I  would  say  that  more  or  less 
of  the  sori  upon  each  frond  are  generally  straight ;  some  are  ham- 
ate, like  a  shepherd's  staff  ;  while  in  other  varieties  they  are  bent 
double  and  become  hippocrepiform  or  almost  round  by  age.  If  we 
ignore  the  genus  Athyrium,  it  is  easy  to  see  how  one  authority  can 
be  justified  for  \AajCttvgfiKx-foetnina  in  Asplenium,  while  another  is 
equally  justifiable  in  calling  it  an  Aspidium.  It  has  seemed  to  me 
that  the  wiser  course  is  to  accept  Athyrium  as  a  valid  genus,  fol- 
lowing the  example  of  Roth  and  Newman  and  Moore,  the  men  who 
have  made  the  closest  study  of  the  genus  and  the  most  detailed  ex- 
amination of  this  particular  species. 

Another  character  of  the  genus  which  is  distinctly  laid  down  in 
Roth's  description  of  it,  is  the  "  laciniate-fimbriate  margin"  of  the 
involucre.  There  is  a  difference  of  opinion  among  authors  as  to 
the  exact  value  of  such  a  character  as  this,  even  when  it  is  constant. 
That  it  is  not  constant,  can  be  certffied  to  by  all  who  have  examined 
any  considerable  number  of  specimens  of  our  American  A.  filix- 
foemina.  The  free  edge  of  the  indusium  is  generally  irregular  and 
somewhat  lacerated,  but  it  could  not  properly  be  called  "  laciniate- 
fimbriate."  But  the  case  is  different  with  Athyrium  cyclosorum. 
Perhaps  no  better  evidence  of  the  validity  of  the  genus  and  of 


—28— 

cvclosorum  as  a  species  could  be  found  than  by  taking  a  vomit; 
frond  of  A.  filix-foemina  rubellum  just  after  the  laciniae  have  un- 
folded in  the  spring,  and  a  frond  of  A. .  cyclosorum  at  the  same 
stage  of  growth.  In  rubellum  the  sori  are  seen  as  distinctly  as 
when  mature,  and  even  at  this  early  period  they  are  beautifully 
hamate,  and  the  outer  edge  of  the  indusium  is  simply  a  little  rag- 
ged. That  is  all,  and  there  is  hardly  enough  of  the  raggedness 
even  to  be  noticeable.  On  the  contrary,  the  sori  on  the  just  open- 
ing fronds  of  A.  cyclosorum  strictum  are  more  often  hippocrepi- 
form  than  hamate,  and  are  sometimes  nearly  umbilicate,  as  if  they 
were  drawn  together  with  a  puckering  string  on  the  side  attached 
to  the  vein  ;  while  the  outer  edge  is  thickly  fringed  with  long  cilia 
plainly  discernible  to  the  naked  eye,  and  which  under  a  magnifier 
are  seen  to  be  many-jointed.  While  these  facts  show  the  nature  of 
the  genus  in  both  cases,  they  also  show  that  the  fringed  indusium 
can  hardly  be  regarded  as  a  generic  distinction,  but  that  in  the  case 
of  A.  cyclosorum  it  forms  a  very  beautiful  instance  of  specific  dif- 
ference. 

If  then  we  allow  that  Athyrium  is  a  valid  genus,  the  question 
naturally  arises,  "  What  species  shall  be  admitted  as  belonging  to 
the  genus?"  I  should  answer  :  all  species  which  conform  to  Roth's 
description  of  the  genus  ;  that  is,  all  species  endowed  with  a  greater 
or  less  number  of  hamate  or  hippocrepiform  sori  and  indusia,  that 
being  the  character  upon  which  Roth  relied  in  distinguishing  the 
genus  from  Polystichum  and  Asplenium.. 

Sir  William  Hooker,  in  his  Species  Filicum,  threw  discredit 
upon  Athyrium  by  including  with  it  as  a  sub-section,  Robert 
Brown's  genus  Allantodia,  which  was  founded  on  species  having 
a  swollen  indusium  and  a  straight,  sausage-shaped  sorus,  as  the 
name  indicates.  He  also  stated  that  "those  who  maintain  the 
genus  (Athyrium}  are  by  no  means  agreed  as  to  the  species  that 
should  be  included  in  it."  That  was  not  a  fair  statement.  If  he 
were  going  to  judge  the  genus  fairly  he  should  have  taken  it  as  its 
author  formed  it  and  not  as  others  had  made  it  by  introducing  char- 
acters which  the  author  did  not  use  and  evidently  did  not  intend 
should  be  used  in  the  genus.  Hooker  himself  was  one  of  the  worst 
of  these,  for  in  his  characterization  of  Athyrium  as  a  sub-genus  of 
Asplenium,  he  says:  "Sori  generally  short;  involucres  lax,  con- 
vex, straight,  or  often  more  or  less  arcuate  and  even  hippocrepi- 
form, sometimes  with  the  lobes  unequal."  He  then  includes  A II- 
antodia  as  a  section  of  Athyrium,  giving  as  its  character,  "  Invo- 


— 29— 

lucres  quite  terete,  very  membranaceous,  tender  and  brittle,  often 
bursting  irregularly."  So  far  as  I  can  judge,  this  is  purely  gratui- 
tous on  Hooker's  part,  and  was  entirely  foreign  to  the  original  idea 
of  Roth.  In  conformity  with  that  idea,  no  species  possessing 
habitually  straight  sori  has  any  place  in  the  genus.  Every  one  of 
the  species  and  varieties  which  he  names  has  more  or  less  arcuate 
sori  ;  and  Hooker's  unfairness  was  accentuated  when  he  placed 
A.  fontamim  under  Eu-Asplenium,  but  described  the  involucres  as 
"very  small,  athytioid"  thus  showing  that  he  recognized  the 
athyrioid  involucre  as  a  valid  type. 

In  conclusion,  there  are  two  or  three  points  which  I  would  like 
to  bring  out  and  make  prominent.  First  is  the  fact  that  every  one 
of  the  species  which  Roth  originally  put  into  his  genus  Athyrium 
was  taken  from  the  Linnaean  genus  Polypodium,  and  was  never 
included  by  Linnaeus  in  the  genus  Asplenium,  in  fact  was  not  con- 
sidered as  having  any  relation  to  Asplenium. 

Secondly,  Presl  in  his  remarks  on  the  tribe  Aspidiae  and  the 
manner  in  which  it  was  broken  up  by  different  authors,  said  : 
"  First  came  Roth,  who  in  the  third  volume  (1800)  of  the  Flora 
Germanica  divided  the  indusiate  Polypodia  of  Linnaeus  into  more 
genera,  viz.  into  Athyrium,  Polystichum,  and  Cyathea.  The  dis- 
tinguished Bernhardi  drove  Athyrium  into  Asplenium  (the  word 
Presl  uses  is  repulsit,  which  is  much  stronger  than  removit  would 
have  been),  accepted  Polystichum,  but  changed  Cyathea  into  Cys- 
topteris,  on  account  of  another  genus  so  called  by  Smith."  So  we 
see  that  until  Bernhardi  "drove"  Athyrium  into  Asplenium, '  the 
species  which  composed  Athyrium  had  no  relation  with  Asplenium 
whatever. 

Thirdly,  Roth  himself,  as  I  have  already  shown,  distinctly  dif- 
ferentiated Athyrium  from  both  Polystichum,  which  was  his  own 
genus,  and  the  Asplenium  of  Linnaeus,  and  asserted  that  the  char- 
acter of  its  sorus  and  indusium  entitled  it  to  be  separated  from  these 
genera  and  to  have  a  genus  of  its  own. 

These  points  seem  to  me  not  only  to  establish  Athyrium  as  a 
\-alid  genus,  but  to  separate  it  wholly  from  any  of  the  other  genera 
with  which  it  has  hitherto  been  associated. 


— 30— 

ON  THE  OCCURRENCE  OF  THE  HART'S=TONQUE 
IN  AHERICA.* 

Bv  WILLIAM  R.  MAXON. 


The  following  account  comprises  chiefly  an  historical  narrative 
of  the  discovery  of  the  American  stations  for  the  Hart's-tongue,  t 
I'hyllitis  Scolopendrium  (L. )  Newm.,  \  with  descriptions  of  habitat, 
especial  mention  being  made  of  the  natural  conditions  which  appear 
to  determine  a  suitable  environment  for  the  fern.  Attention  has 
been  given  also  to  the  question  of  the  fern's  former  distribution, 
and  an  attempt  made  to  determine  the  principal  causes  that  have 
operated  to  effect  the  peculiarly  limited  distribution  of  the  present. 

I  shall  discuss  the  American  stations  substantially  in  the  order 
of  their  discovery.  Considerable  care  has  been  exercised  in  sup- 
plying in  full  and  verifying  all  available  references  bearing  directly 
upon  the  subject.  The  central  New  York  stations  are  the  only 
ones  writh  which  I  am  personally  familiar.  These  I  have  frequently 
visited  at  various  times  from  1895  to  1898,  while  living  at  Oneida 
and  Syracuse,  N.  Y.  To  the  several  correspondents,  mentioned 
later,  who  have  generously  furnished  data,  largely  the  result  of 
personal  observation,  I  would  extend  my  sincere  thanks.  || 

CENTRAL  NEW  YORK  STATIONS. 

In  central  New  York  the  Hart's-tongue  has  been  found  grow- 
ing in  four  separate  localities  :  (a)  Geddes  ;  (b)  Chittenango  Falls  ; 
(c)  Jamesville  ;  (d)  Perryville.  The  range  of  territory  covered  is 
comparatively  small,  and  the  stations  are  separated  by  short  dis- 
tances only.  Each  station  is  entirely  distinct,  however,  and  scat- 
tering plants  seem  not  to  occur  in  the  intervening-  territority. 

*  Published  by  permission  of  the  Secretary  of  the  Smithsonian  Institution. 
tThe  term  is  applied  here  only  to  the  species  Scolopendrium. 
\  Principal  synonomy  : 

Asplenium  Scolopendrium    L.  Sp.  PI.  1078.     1753. 

Scolopendrium  vulgare  J.  E.  Smith,  in  Mem.  Acad.  Tur  5:  421.     1793. 

Scolopendrium  officinarum   Sw.   in   Schrad.  Journ.   Hot.    2-  Part    2,  61. 

(1800)  1801. 

Phyllitis  Scolopendrium  (L.)  Newm.  Hist.  Brit.  Ferns,  Ed  2.  271.     1854. 
Scolopendrium  Scolopendrium  (i,.)  Karst.  Deutsch.  Fl.  Ed.  i.  278.  1880-83. 
II  A  large  part  of  the  present  paper  is  extracted  from  aa  unpublished  thesis 
by  the  writer  presented  to  the  faculty  of  Syracuse  University,  June,  1898,  en- 
titled :  "  A  Contribution  to  the  Biology  of  the  Hart's-tongue  Fern." 


a.  The  Geddes  Station.  The  Hart's-tongne  was  first  discov- 
ered in  America  by  Frederick  Pursh,  on  July  20,  1807,  near  the 
place  now  known  as  Split  Rock,  a  small  suburb  of  Syracuse,  about 
five  miles  west.  Apparently  the  first  mention  in  literature  of  this, 
the  earliest  American  station,  is  contained  in  Pursh's  Mora,*  in 
which  the  following  statement  occurs:  "In  shady  woods,  among 
loose  rocks  in  the  western  part  of  New  York,  near  Onondago,  on 
the  plantation  of  J.  Geddis,  Esq.  Perennial.  July.  v.  v.  This 
species  I  have  seen  in  no  other  place  but  that  here  mentioned,  nei- 
ther have  I  any  information  of  its  having  been  found  in  any  other 
part  of  North  America."  Subsequent  search  failed  to  again  dis- 
cover the  fern  in  this  locality,  though  it  was  found  at  Chittenango 
Falls  (about  1830)  and  in  the  Jamesville  vicinity  (1857).  The  latter 
discovery  doubtless  led  to  the  renewed  attempts  to  find  the  plants 
at  Geddes. 

At  the  suggestion  of  Dr.  Asa  Gray,  Mr.  J.  A.  Paine,  in  June, 
1866,  visited  the  locality  for  the  purpose  of  verifying  Pursh's  origi- 
nal station.  In  his  account  of  the  trip  f  he  says,  after  describing  a 
fruitless  search  :  "  Hon.  George  Geddes,  son  of  the  J.  Geddes, 
Ksq.,  referred  to  by  Pursh,  was  then  appealed  to  for  information  in 
general  respecting  this  fern  or  its  earliest  station,  and  he  readily 
cleared  up  the  whole  mystery.  The  place  where  it  was  discovered, 
he  said,  was  nearly  five  miles  west  of  Syracuse  and  half  a  mile 
south  of  his  father's  house  ;  on  the  single  point  of  its  being  on  his 
father's  farm  Pursh  must  have  erred  ;  but  it  was  nearby  along  a 
high  ledge  and  about  a  celebrated  sulphur  spring."  Paine  did  not 
succeed  in  rediscovering  the  fern  at  Geddes,  and  it  was  generally 
supposed  that  it  no  longer  persisted  there,  until  in  1879  (September 
3oth )  it  was  rediscovered  in  fair  quantity  upon  the  Geddes  property 
by  members  of  the  Syracuse  Botany  Club.  ;  The  ferns  continued 
to  grow  thriftily  until  the  summer  of  1895,  when  the  Solvay  Soda 
Ash  concern  blasted  out  the  rocks  which  had  so  long  served  as  a 
shelter.  It  is  extremely  unlikely  that  a  single  plant  has  survived. 

It  seems  strange  that  up  to  1879  no  one  should  have  rediscov- 
ered the  fern  here.  It  may  be  of  interest  to  note  that  the  supposi- 
tion of  Mr.  George  Geddes  and  Paine  that  Pursh  was  in  error  in 
his  statement  of  its  occurrence  upon  the  farm  is  not  borne  out  by 

*  Pursh,  Fl.  Am.  Sept.  2:  667.     1814. 

t  Am.  Journ.  Sci.  &  Arts,  II.  42:  282.     1866. 

J  Hull.  Torr.  Bot.  Club.  Q:  345-7.     1879. 


—32— 

the  facts  of  its  discovery  in  1879.  Mrs.  L.  L.  Goodrich,  of  Syra- 
cuse, a  relative  of  the  Geddes  family,  and  a  member  of  the  discov- 
ering party,  has  stated  positively  to  me  that  the  fern  was  upon  that 
occasion  found  in  considerable  abundance  well  within  the  limits  of 
the  original  "Geddes  plantations." 

Pursh  further  says  of  this  station  :  *  "  Mr  Geddes  brought  me 
to  a  deep  valley  about  i.  m.  from  his  house,  where  we  ascended 
a  steep  very  rocky  hill  ;  here  large  masses  of  rock  seem  to  be  piled 
up  or  turned  over  one  &  another  in  such  a  confused  manner,  that 
it  has  left  large  chasms  between  them,  which  sometimes  appear 
like  caves  :  as  it  has  a  north  aspect  &  overshadet  with  trees,  all  the 
rocks  are  covered  with  moss  and  vegetables  :  *  *  *  &  what  I 
thought  most  of  Asplenium  Scolopendriuni — this  fern  which  I  dont 
find  mentioned  by  any  one  to  grow  in  America  I  allways  had  a 
notion  to  be  here  ;  &  indeed  I  was  quit  enjoyed  to  find  my  preju- 
dice so  well  founded  in  truth.  It  appears  Jo  be  the  same  as  the 
european  only  smaler ;  query  ?  is  the  european  auriculated  at  the 
base  like  this  species?" 

Split  Rock  is  a  limestone  formation,  consisting  mostly  of  the 
Lower  Helderberg,  which  extends  to  a  depth  of  approximately  125 
feet,  with  the  so-called  Oriskany  sandstone  interposing  in  a  very 
thin  sheet,  from  one  to  five  inches  thick,  between  it  and  the  Cor- 
niferous  of  the  Upper  Helderberg  (12  to  14  feet  thick)  which  caps 
the  plateau.  Dr.  Underwood  has  several  times  t  called  attention  to 
the  fact  that  in  central  New  York  at  least,  this  fern  is  found  only  at 
the  outcrop  of  the  Corniferous.  It  was,  before  the  inroad  of  the 
quarryman,  a  steep  ledge  about,  150  feet  high  and  half  a  mile  long, 
somewhat  semicircular  in  general  outline.  At  the  base  of  the  cliff 
was  a  brook,  with  a  sulphur  spring  upon  its  bank.  It  was  at  some 
little  distance  from  this  spring,  I  am  told,  that  the  plants  grew, 
somewhat  sheltered  by  rocks  and  trees,  but  to  a  large  degree  ex- 
posed to  the  sweep  of  the  cold  northwest  winds. 

b.  The  Chittenango  Falls  Station.  The  second  place  in  Amer- 
ica at  which  this  fern  is  known  positively  to  have  been  found,  is 
Chittenango  Falls,  in  -Madison  county,  where  Mr.  William  Cooper 
found  it  about  1830.  From  1830  to  1857  it  was  the  only  American 

*  Frederick  Pursh  :  A  Journal  of  a  Botanical  Excursion  in  the  Northern 
Parts  of  the  States  of  Pennsylvania  and  New  York  during  the  year  1807.  63-64. 
Edited  by  T.  P.  James,  aud  issued  1869.  pp.  87. 

t  Underwood  :  Our  Native  Ferns,  Ed  5,  p.  6.  1896.  Also  in  Britton  &  Brown, 
Illus.  Flora  North.  U.  S.  and  Can.  \:  21.  1896. 


station  definitely  known,  *  and  perhaps  on  this  account  Pursh's  dis- 
covery has  often  been  incorrectly  assigned  to  Chittenango  Falls. 

The  Chittenango  creek,  flowing  northward  toward  Oneida  lake, 
here  takes  a  double  plunge  of  over  one  hundred  feet,  and  has  worn 
through  the  limestone  a  rough  gorge  of  that  depth  and  more.  The 
sides  are  extremely  steep,  but  debris  and  soils  have  so  accumulated 
at  the  base  that  the  unbroken  ledges  loom  up  only  along  the  top  of 
the  gorge.  It  is  just  out  from  under  these  overhanging  cliffs  and 
among  the  broken  fallen  fragments  of  limestone  ( mostly  Cornifer- 
ous)  on  the  left  bank,  that  the  Hart's-tongue  grows,  perhaps 
thirty-five  or  forty  feet  above  the  level  of  the  stream,  and  three  or 
four  rods  distant.  The  soil  is  moderately  moist,  but  light,  yielding, 
and  very  rich  in  leaf  mould.  Here,  scattered  along  the  steep  bank 
for  a  distance  of  nearly  a  quarter  of  a  mile  from  the  falls,  the  ferns 
grow  in  the  fairly  dense  shade  of  second-growth  maples,  beeches, 
birches  and  elms.  Among  its  companion  plants  are  Pellaea  Stel- 
/eri,  Asplenintn  Kitta-nmraria,  Cystopteris  bulbifera,  and  Dryop- 
teris  Goldieana. 

c.  The  Jamesville  Locality.  The  Hart's-tongue  grows  abund- 
antly in  a  number  of  places  in  the  immediate  vicinity  of  Jamesville, 
the  distance  included  being  ten  to  thirteen  miles  west  of  Chittenango 
Falls,  and  four  to  nine  miles  southeast  of  Syracuse.  Exceptionally- 
fine  plants  grow  in  this  locality,  and  almost  the  same  conditions 
obtain  in  each  of  the  stations,  which  are  (i)  Hoivletf  s  Gorge; 
(2)  Little  (or  Green)  Lake;  (3)  Green  Pond;  (4)  Rock  Gorge. 

(1)  The  Hewlett"  s  Gorge  Station.     In  March,  1866,  Mr.  Lewis 
Foote,  of  Detroit,  Mich.,  found  the  fern  growing  plentifully  in  a 
deep  ravine  of  Butternut  creek,  five  miles  southeast  of  Syracuse, 
upon   the   line  of   the   Syracuse   &    Binghamton    Railway,  f     This 
ravine  is  commonly  known  as  Howlett's  Gorge.     It  is  deep  and 
rocky,  especially  rough  on  the  left   or   northern   side  where   the 
Hart's-tongue  grows,  though  more  open  on    the    opposite    side. 
The  fern,  well  shaded,  once  grew  here  very  plentifully,  but  it  has 
been  largely  rooted  out. 

(2)  Little  (or  Green]  Lake.     In  September  of  the  same  year, 
Mr.  J.  A.  Paine  visited  the  locality  of  Jamesville  and  extended  the 
known  range  nearly  to  its  present  limits.     He  detected  the  fern  on 


*  Asa  Gray  in  Am.  Journ.  Sci.  &  Arts,  II.  41:417.     1866.     I  have  been  unable 
to  find  any  earlier  reference  to  Cooper's  discovery. 

t  Asa  Gray,  in  Am.  Journ.  Sci.  &  Arts,  II.  41 :  41?-     l866- 


—34  — 

the  shaded  talus  of  cliffs  which  nearly  surround  Little  Lake.  Little 
Lake  is  situated  approximately  a  mile  south  of  Hewlett's  Gorge, 
and  a  mile  west  of  Jamesville.  Paine  aptly  describes  it  as  "a  deep 
depression  in  the  surface,  walled  in  on  all  sides  but  one  with  rocks 
at  least  100  feet  high,  and  one-fourth  of  a  mile  across  from  side  to 
side."  *  The  open  side  is  the  eastern,  t  The  plant  was  formerly 
very  abundant  on  the  talus  at  the  south  of  the  lake,  but  scarcely  a 
half  dozen  plants  may  be  found  now,  owing  to  the  greed  of  picnick- 
ers. From  thirty  to  fifty  rods  to  the  north  of  the  lake  the  plant 
grows  thriftily  in  at  least  three  different  places  along  the  sides  of 
two  wooded  ravines  which  occur  together. 

(3)  Green  Pond.  Continuing  his  search,  Mr.  Paine  gave  at- 
tention to  the  other  pit-hole  lakes  of  the  vicinity,  and  found  Phyllitis 
growing  at  Green  Pond.  White  Lake  and  Green  Pond  lie  near 
each  other,  a  mile  and  a  half  east  of  Jamesville,  at  the  base  of  a 
ledge  of  limestone  from  100  to  200  feet  high.  This  ledge  is  a  con- 
tinuation eastward  of  the  steep  escarpment  which  forms  the  south- 
ern cliff  of  Rock  Gorge,  lying  about  a  mile  northwest  of  Little 
Lake.  As  stated,  it  extends  eastward,  and  transecting  the  north- 
erly-trending Butternut  valley,  runs  a  half  dozen  miles  farther,  inci- 
dentally giving  rise,  at  a  given  point  along  its  base,  to  Green  Pond. 

Green  Pond  (also  called  Scolopendrium  Lake)  is  similar  to 
Little  Lake  in  lying  like  a  sheltered  harbor  far  within  the  irregular 
outline  of  the  surrounding  cliffs.  It  is,  however,  at  least  a  third  of 
a  mile  broad.  The  banks  are  exceedingly  rough  and  strewn  with 
fragments  broken  from  the  towering  limestone  cliffs.  %  The  cliffs 
have  extensive  tali,  and  it  is  the  continuous  steep  talus  of  the  great 
U-shaped  cliff  which  forms  the  shore  of  the  lake.  The  fern  grows 
pretty  well  up  on  the  sides,  among  the  fragments  of  Corniferous 
limestone,  on  both  sides  at  the  base  of  the  U.  The  plants  from  the 
cleared  (eastern)  portion  have  mostly  become  of  small  size  and 
winter-kill  badly,  owing  doubtless  to  the  comparatively  recent 
removal  of  the  forest,  which  occasions  a  lack  of  protection  in  win- 
ter and  summer  alike.  On  the  western  slope,  as  yet  wooded,  the 
plants  grow  to  good  size. 


*  Amer.  Journ.  Sci.  &  Arts,  II.  42:  281.     1866. 

fFor  a  complete  description  of  this  remarkable  lake  and  vicinity,  see  the 
article  by  Prof.  E.  C.  Quereau,  entitled  Topog.  and  Hist.  Jamesville  Lake,  N.Y.. 
111  Bull.  Geolog.  Soc.  Am.  9:  173-182.  1898. 

t  See  article  by  the  author,  Fern  Bull.  7:1.     1899. 


—35— 

(4)  Rock  Gorge  Station.  Rock  Gorge,  lying  about  a  mile 
northwest  of  Little  Lake,  is  one  of  the  more  northerly  transverse 
valleys  connecting  the  Onondaga  and  Butternut  valleys.  It  runs 
east  and  west,  trending  slightly  to  the  southeast,  and  is  utilized  by 
the  Syracuse  &  Binghamton  Railway.  At  a  point  about  "  midway 
its  length,  and  on  the  south  side  of  the  gorge,  the  wall  is  cut  back 
in  the  form  of  an  ampitheatre  which  is  semicircular  in  outline  and 
about  125  feet  deep  by  250  feet  wide.  The  walls  are  nearly  per- 
pendicular, with  their  bases  concealed  by  recent  talus  accumula- 
tion." *  In  this  recess,  about  40  feet  from  the  top  of  the  cliff,  and 
among  the  loose  fragments,  grow  about  125  extremely  fine  plants 
of  nyllitis.  The  slope  is  rather  steep,  but  the  plants  grow  thriftily 
in  the  scattering  second  growth  of  maple  and  basswood,  shaded  by 
the  cliff  wall,  which  serves  also  as  a  considerable  protection  in 
winter.  ^ 

A  small  number  of  plants  have  also  been  observed  recently  ( in 
May,  1899)  to  grow  in  a  small  depression  some  40  rods  to  the  west- 
ward and  back  from  the  amphitheatre,  by  Mr.  Homer  D.  House. 

Mr.  Paine,  at  the  time  of  his  vtsit  to  Jamesville  in  1866, 
gave  a  great  deal  of  attention  to  the  general  contour  of  the 
locality.  He  remarks f  that  "these  'highlands'  before  they  were 
cleared  and  burned  over,  formed  the  very  kind  of  locality  where 
our  rare  fern  delights  to  dwell,  possessing  all  the  conditions  of 
loose  limestones,  rich  mould,  moisture  and  shade  ;  and  no  doubt 
their  rocky  steeps  formerly  abounded  with  it.  This  presumption 
is  confirmed  by  the  fact  that  on  a  particular  part  of  the  range,  where 
the  fire  and  clearing  ceased  and  the  undisturbed  forest  began,  just 
there  was  Scolopendrium  found  growing  in  its  greatest  luxuriance 
and  scattered  along  the  bank  for  a  fourth  of  a  mile  or  so,  as  far  as 
covered  by  rocks." 

d.  The  Perryville  Station.  An  additional  central  New  York 
station  for  the  Hart's-tongue  was  discovered  in  July,  1898,  at 
Perryville  Falls,  Perryville,  by  Miss  Murray  Ledyard,  of  Cazenovia, 
N.  Y.  A  small  stream,  the  Canaseraga  creek,  here  falls  fully  a 
hundred  feet,  near  the  quarries  at  the  railway  station,  and  runs 
helter-skelter  through  the  narrow  wooded  ravine  below.  As  at 


*See  Prof.   Quereau's  paper  previously  mentioned.     He   finds  conclusive 
that  this  amphitheatre  was  once  a  waterfall, 
t  Amer.  Journ.  Sci.  &  Arts,  II.  42:  281.     1866. 


-36- 

Chittenango  Falls,  the  fern  grows  only  upon  the  western  side  of  the 
gorge;  and  the  two  stations  are  otherwise  very  similar.  Mrs.  James 
R.  Parsons,  one  of  the  discovering  part}',  thus  writes  of  its  occur- 
rence :*  "  The  plants  were  *  *growing  in  a  partial  open- 
ing among  the  maples,  basswoods  and  beeches  on  a  steep  slope 
covered  with  fragments  of  limestone,  some  30  or  40  feet  from  the 
base  of  the  cliff.  We  must  have  found  anywhere  from  20  to  30 
plants  within  a  radius  of  as  many  feet."  The  fact  of  its  discovery 
here,  in  a  favorite  botanizing  field,  may  indicate  a  recent  origin  of 
this  particular  station.  It  may  have  arisen  from  the  Chittenango 
station,  which  is  less  than  three  miles  distant. 

In  general,  regarding  its  central  New  York  distribution  :  It  is 
extremely  likely  that  the  fern  has  by  the  natural  clearing  of  the 
country  been  in  some  measure  exterminated  ;  but  it  is  a  fact,  nev- 
ertheless, that  it  is  only  in  the  more  rugged  situations  of  the  un- 
cleared land  that  it  usually  grows.  It  stands  rather  as  a  remaining 
type  of  boreal  vegetation,  persisting  only  in  such  places  as  are  well 
suited  to  it.  It  prefers  rough,  shaded  tali  in  broken  country,  where 
extreme  drought  can  never  affect  it ;  where  it  is  subjected  to  a  uni- 
formly cool  temperature,  and  protected  also  by  considerable  shade. 
In  such  a  situation  the  fern  now  thrives,  and  doubtless  will  so  con- 
tinue unless .  rooted  out  by  reckless  collectors.  It  ought  even  to 
become  settled  in  many  additional  stations  in  the  general  locality. 

CANADIAN  STATIONS. 

(a)  The  Owen  Sound  Locality.  Owen  Sound  is  a  port  on  the 
(Georgian  Bay,  the  great  eastern  arm  of  Lake  Huron.  The  town  is 
nearly  on  the  lake  level,  the  rise  being  perhaps  fifty  feet  to  the 
mile  ;  while  both  east  and  west  are  cliffs— Clinton  on  the  west  side, 
Medina  on  the  east— which  form  the  sides  of  the  valley.  The  rock 
is  covered  with  soil  of  varying  shallowness,  and  forested  with 
maple,  spruce,  hemlock,  and  birch.  The  country  all  about  is  very 
rocky,  and  doubtless  contains  more  stations  than  mentioned  here. 

The  first  discovery  of  the  fern  at  Owen  Sound  was  by  Prof. 
William  Hinks,  of  Toronto,  in  1857.  He  found  it  growing  plenti- 
fully around  the  falls  of  a  stream  emptying  into  the  Sound,  t  This 
stream  is  the  Sydenham  river,  and  the  falls,  which  are  situated  over 
two  miles  south  of  the  town,  are  known  as  ' '  Sydenham ' '  or 

*  Fern  Bull.  4:  74.     iSgS. 

t  J.  A.  Paine,  in  Am.  Journ.  Sci.  &  Arts,  II.  42:  2Si.     iS66. 


—37— 

"  Inglis"  Falls.  Below  the  falls  occur  the  plants  of  Phyllitis,  in  a 
"  fairly  heavy  \vood  in  the  valley  and  on  the  sloping  sides  of  the 
deep  chasm  through  which  the  Sydenham  runs  after  its  fall.  The 
chasm  is  about  a  fourth  of  a  mile  wide,  or  less,  its  sides  strewn 
with  boulders  and  fragments  of  limestone  partially  buried  in  debris. 
Here  the  individuals  were  seldom  large,  i.  e.  seldom  over  8  inches 
long.*  The  fern  is  reckoned  "abundant  on  limestone  debris 
under  cliffs  at  Sydenham  Falls  and  other  localities  around  Owen 
Sound  (Mrs.  Roy)."  t 

It  has  also  been  found  close  by  "growing  in  a  maple  wood, 
where  the  rock  is  c  lose  to  the  surface  and  shows  cracks  of  width 
varying  from  two  inches  to  two  feet.  It  grows  [here]  generally  in 
the  cracks,  but  also  on  the  level  ground,  doubtless  always  where 
the  soil  is  shallow.  Companion  plants  are  Dryopteris  marginalis, 
I'olystichum  lonchitis,  Camptosorus,  and  Aspleniiini  viride.  % 

Phyllitis  occurs  also  in  a  wild  situation  some  twelve  miles  to 
the  northwest  of  the  village.  Here,  too,  the  soil  is  not  deep,  and 
the  fern  grows  rather  thriftily  in  the  dark,  moist,  rocky  woods, 
where  the  limestone  ( Clinton )  comes  close  to  the  surface.  In  such 
places  it  grows  on  small  hummocks  slightly  raised  above,  and  so 
drier  than  the  surroundings.  -' 

The  finest  plants  are  found  at  a  point  about  one  mile  northwest 
of  the  town,  upon  the  loose  limestone  debris  fallen  from  a  bluff  40 
feet  high.  The  soil  is  very  light,  porous,  and  rather  dry,  and  can- 
not for  any  length  of  time  retain  ijjoisture.  But  the  situation  seems 
especially  favorable,  and  fronds  grow  from  eighteen  to  twenty- 
three  inches  long.  :;~ 

1'hyllitis  grows  about  Owen  Sound  very  much  as  in  New  York, 
in  loose  limestone  debris,  and  additionally,  in  narrow  limestone 
crevices  and  raised  on  hummocks  in  moist,  rocky,  upland  woods — 
the  sort  of  situations  especially  claimed  for  it  formerly  in  central 
New  York  by  Paine.  Mr.  Jenkins  obtains  the  best  developed 
specimens  from  the  station  a  mile  northwest  of  the  town,  where 
the  soil  is  the  driest  and  most  porous  of  any  of  the  stations.  Fur- 

*  From  correspondence,  Prof.  W.  II.  Jenkins,  Owen  Sound,  Ontario,  Canada. 

tMacoun,  Cat.  Can.  PI.  Part  V.,  268.     1890. 

%  From  correspondence  with  Mr.  W.  K.  Saunders,  I,ondon,  Ontario. 


200775 


Iher,  he  states  that  the  summer  of  1897  being  a  "wet  one,  the 
t'ronds  were  somewhat  shorter  and  narrower  than  in  previous 
years."  The  habitat  of  the  fern  is,  in  each  case,  cool,  well  shaded, 
and  invariably  upon  limestone.  But  Mr.  Jenkins  remarks  further 
that  while  each  is  "moist  so  far  as  atmosphere  is  concerned,"  it 
appears  that  "  moisture  as  a  soil  constituant  is  not  of  prime  import- 
ance to  large  growth." 

(b)  The  Durham  Station.  The  fern  was  found  in  1883,  "on 
Guelph  dolomites,  Little  Sau  river,  at  Durham,  Gray  county,  On- 
tario," by  Dr.  H.  M.  Ami,  of  the  Canadian  Geological  Survey/"' 
who  has  written  me  as  follows:  "The  specimens  obtained  were 
growing  in  the  narrow  crevices  of  the  cream-colored  dolomites  of 
the  Gnelph  formation  (Silurian).  They  appeared  to  be  somewhat 
depauperate  forms,  still  sufficiently  alive  and  vigorous  to  warrant 
the  expectation  that  they  would  survive  under  the  existing  environ- 
ment. There  was  very  little  earth  where  they  were  growing,  the 
rock  everywhere  being  practically  bare  or  destitute  of  earth  or 
drift."  Durham  is  about  20  miles  south  of  Owen  Sound  in  a  very 
wild  country,  little  explored  botanically.  Guelph  dolomite  is  a 
good  half  magnesium  carbonate. 

,  (c)  The  Col  ling  wood  Station.  1111898.  Prof.  Jenkins  wrote  me 
of  the  supposed  occurrence  of  Phyllitis  at  Collingwood,  Ontario. 
Lately  I  have  learned  through  Mrs.  E.  G.  Britton  of  its  rediscovery 
by  Mr.  B.  B.  Osier,  of  Toronto,  a  member  of  this  Chapter.  Mr. 
Osier  has  kindly  communicated  tnost  of  the  following  data,  which 
are  set  down  largely  verbatim  :  The  location  is  lot  XI.  in  the  3d 
concession  of.  the  township  of  Collingwood,  Grey  county,  Ontario. 
It  is  seven  miles  in  a  westerly  direction  from  the  town  of  Colling- 
wood, which  is  on  the  Georgian  Bay,  and  about  50  miles  east  of 
Owen  Sound.  The  land  is  about  1500  feet  above  sea  level  and  of 
the  Upper  Silurian  formation,  a  limestone  of  sufficient  purity  to  be 
burned  in  neighboring  kilns.  The  forest  is  ordinary  Ontario 
growth  of  hard  maple,  mountain  maple,  basswood,  elm,  ash,  beech 
and  iron-wood,  with  more  or  less  cedar  and  butternut.  The  soil  is 
a  rich  clay  loam,  with  a  great  deal  of  leaf  mould.  The  rock  on 
which  the  ferns  grow  is  full  of  seams  and  crevices,  which,  together 
with  the  absence  of  quick  evaporation  (due  to  the  dense  shade), 
generally  gives  ample  moisture.  The  region  is  essentially  a  plat- 

*  Macouii,  Cat.  Can.  PI.  Part  V.  268.     1890. 


—39— 

eau,  cut  into  by  several  streams  which  have  made  valleys  and 
gorges  trending  mostly  to  the  east.  The  most  northerly  of  these  is 
the  Silver.  The  Silver  rises  in  a  group  of  springs  flowing  perhaps 
20,000  gallons  per  hour.  The  ground  is  moist,  the  shade  dense, 
and  the  ground  largely  boulder  limestone.  The  ferns  are  scattered 
over  approximately  two  acres  immediately  surrounding  the  springs. 
The  point  of  first  discovery  lies  some  120  rods  to  the  northeast  of 
tli is  point,  upon  the  rocky  tali  and  slopes  above  Kennedy  creek. 
A  few  plants  occur  also  near  the  banks  of  the  Silver,  about  a  mile 
from  its  source.  The  valley  of  the  Pretty  lies  about  two  miles  to 
the  south  of  the  Silver.  The  fern  occurs  here  in  some  abundance 
along  the  rocky  slopes  of  the  valley,  though  the  land  is  higher  and 
drier,  and  the  shade  not  so  dense.  Many  granite  and  schist  bould- 
ers occur  here,  but  do  not  carry  Hart's-tongue,  the  fern  being 
always  rooted  in  the  limestone  crevices. 

Mr.  Os-ler  is  of  the  opinion  that  the  fern  is  also  to  be  found  in 
the  valley  of  the  Pine  and  the  Mad,  the  formation  and  aspect  being 
similar.  The  Holly  fern  always  occurs  with  the  Hart's-tongue  at 
these  stations,  and  near  the  springs  the  Walking-leaf  in  consider- 
able abundance.  Mr.  Osier  has  kindly  furnished  a  series  of  photo- 
graphs of  the  ferns  in  their  native  environment.  In  a  later  letter, 
attention  is  called  to  the  fact  of  the  fern's  destruction  in  quantity 
by  young  cattle. 

(d)  The  Woodstock  Station.  The  Hart's-tongue  was  discov- 
ered near  Woodstock,  N.  B.,  in  the  late  fall  of  1882  by  James  Sut- 
ton,  a  gardener  in  the  employ  of  Mrs.  Charles  Connell,  of  Wood- 
stock. During  a  visit  to  this  village  the  next  September,  the  atten- 
tion of  the  late  Peter  Jack,  of  Halifax,  N.  S.,  was  directed  to  the 
plant,  a  single  rather  undersized  specimen,  which  had  been  pre- 
served in  Mrs.  Connell's  greenhouse.  Mr.  Jack  immediately  rec- 
ognized it  as  "  Scolopendrium  vulgare."  *  From  Mrs.  Dibblee,  nee 
Connell,  of  Woodstock,  and  Mr.  G.  U.  Hay,  of  St.  John,  N.  B.,  I 
have  learned  that  it  was  collected  some  six  miles  to  the  westward  of 
Woodstock,  upon  the  "Richmond  Road,"  near  the  Meduxnakik  river 
where  it  was  open  to  the  northwest.  Mrs.  Dibblee  has  stated  also 
that  the  station  ' '  has  been  all  burnt  over,  ploughed  up,  and  is  now 
a  fine  farm,"  and  that  the  fern  grew  upon  what  is  commonly  called 


*  See  Notice  of  New  and  Rare  Plants  by  George  I,awson,  in  Proc.  and  Trans. 
Nov.  Scot.  lust.  Nat.  Sci.  Q.  71-72.  1883-6 ;  also  a  paper  by  G.  U.  Hay,  entitled 
Potai4y  of  the  I'pper  St.  Joint,  in  Bull.  Nat.  Hist.  Soc.  N.  Br.  No.  2,  pp.  31  and  37. 


—4o— 

there  the  shale  land  which  supports  a  good  growth  of  elms,  butter- 
nuts and  ash. 

Mr.  Jack  visited  the  station  at  the  time,  but  without  finding 
further  plants.  After  his  return  to  Halifax,  the  gardener  upon  dili- 
gent search  secured  a  number  of  additional  plants,  four  of  which 
were  forwarded  to  Mr.  Jack,  who  later  presented  one  to  the  Nat- 
ural History  Society  of  New  Brunswick,  and  a  frond  to  the  Nova 
Scotia  Institute  of  Natural  Sciences.  The  Eaton  Herbarium  con- 
tains also  two  fronds  of  the  var.  marginatum,  collected  by  Sutton 
in  July,  1885.  Mrs.  Dibblee  states  that  the  fern  was  brought  in 
several  times  from  the  same  place.  Mr.  Hay  adds  that  quite  a 
number  of  them  still  thrive  in  the  conservatory  and  upon  the  rock- 
ery at  her  home,  but  that  plants  presented  to  him  at  various  times 
have  not  survived. 

Mr.  John  Macoun  has  (in  correspondence)  suggested  the  pos- 
sibility of  the  fern's  having  become  first  established  as  an  escape, 
but  a  thorough  knowledge  of  the  facts  attending  its  discovery  has 
convinced  both  Mr.  Hay  and  myself  that  it  was  undoubtedly  native. 
In  fact,  Mr.  Sutton  has  stated  that  he  once  found  the  fern  in  a 
ravine  farther  in  the  woods,  about  eight  miles  from  town,  but  that 
he  has  not  again  seen  it  there.  Mr.  Hay  has  long  intended  a  sys- 
tematic search  for  the  fern  about  Woodstock.  Such  a  search  would 
probably  result  in  its  discovery  somewhere  in  the  general  vicinity. 

THE  TENNESSEE  STATIONS.  * 

The  Hart's-tongue  has  been  found  in  two  localities  in  Tenn- 
essee, viz.,  near  Post  Oak  Springs,  and  at  South  Pittsburg,  of  which 
the  latter  only  has  been  known  in  literature. 

a.  The  Post  Oak  Springs  Station.  In  1849  ^r-  A.Gattinger  de- 
tected the  fern  a  short  distance  west  of  the  village  of  Post  Oak 
Springs,  in  Roane  county.  About  one  mile  southwest  of  the  vil- 
lage occurs  a  pool  in  an  open  cave  in  the  front  ( southeastern )  edge 
of  the  hill,  and  from  this  issues  a  small  stream  to  the  eastward. 
Over  the  top  of  the  hill,  i.  e.,  on  the  northern  side,  and  distant 
about  a  half  inile,  occurs  another  open  cave,  called  the  "  dry 
cave,"  about  the  mouth  of  which  a  few  plants  were  found  by  Dr. 
Gattinger.  This  cave  is  about  one  mile  directly  west  of  the  village. 

*  In  his  Tennessee  Flora  ( 1887)  p.  102,  Dr.  Gattinger  refers  the  fern  to  "  New 
Pittsburg,"  a  mistake  for  South  Pittsburg,  and  adds  "not  found  [there]  by 
myself."  The  previous  station  (Post  Oak  Springs)  concerning  which  he  has 
lately  written  me,  was  unfortunately  overlooked. 


According  to  the  Kingston  Folio  of  the  U.  S.  Geological  Sur- 
vey Atlas,  the  immediate  rock  formation  is  the  Knox  Dolomite,  or 
Magnesium  Limestone  of  the  Lower  Silurian.  But  Dr.  Gattinger 
avers  that  the  Chattanooga  Black  Shale  (Devonian)  and  the  Fort 
Payne  Chert  (Lower  Carboniferous)  both  crop  out  here.  The  fern 
probably  occurs  in  the  latter.  *  Immediately  above  is  the  Bangor 
Limestone,  which  supports  the  fern  at  South  Pittsburg. 

b.  The  Smith  Pittsburg  Station.  The  Hart's-tongue  was  discov- 
ered growing  in  a  deep  sink-hole  near  South  Pittsburg,  by  Major 
Cheathem,  in  1879.  t  South  Pittsburg  is  a  town  on  the  Tennessee 
river,  about  three  miles  north  of  the  Alabama  boundary.  Some 
two  miles  southwest  of  the  town,  two  spurs  of  the  Cumberland 
mountains,  extending  southeast  into  the  level  plain  of  the  river, 
form  a  narrow  valley  or  "cove,"  as  they  say  in  Tennessee.  To 
reach  the  sink-hole,  follow  the  cove  a  half  mile,  or  until  half  way  up 
the  mountain.  Sixty  feet  to  the  left  of  this  narrow  valley  and  about 
sixty  feet  above,  there  is  an  irregular  fissnre  in  the  Mountain  lime- 
stone, sixty  feet  long  by  twenty  to  forty  feet  wide  and  ninety-two 
feet  deep.  Upon  examination,  a  good-sized  spring  is  found  to 
issue  from  a  cave  not  more  than  twenty  yards  farther  up  the  hill. 
This  spring,  tumbling  perpendicularly  into  the  hole,  strikes  a  pro- 
jecting ledge  some  forty  feet  below.  The  water  splashing  from 
tth  ledge  has  worn  a  deep  depression  in  the  opposite  side,  and  it 
is  chiefly  upon  this  slope  that  the  fern  grows.  The  area  covered  by 
the  ferns  is  not  over  200  square  feet,  and  contained  (in  1898)  about 
1 10  mature  plants.  A  few  are  variously  distributed  along  the  sides 
and  edges  of  the  chasm,  with  Asplenium  parvulum.  The  soil  is  a 
sticky,  light-colored  clay,  formed  from  the  disintegrated  shales  of 
the  upper  mountain.  The  ferns  are  found  mostly  about  fifty  feet 
below  the  surface,  and  are  so  sheltered  that  some  of  them  the 
direct  sunlight  never  reaches,  and  can  possibly  reach  none  of  them 
longer  than  two  hours  a  day.  There  is  little  variation  in  tempera- 
ture, naturally,  and  they  are  always  dampened  by  the  spray  of  the 
falling  water.  After  its  first  drop  of  forty  feet,  the  water  trickles  in 
small  streamlets  over  the  rock  walls  and  fragments  the  remaining 
fifty-two  feet,  to  disappear  in  a  narrow  fissure  at  the  bottom  of  the 


*  Dr.  Gattinger  has  kindly  furnished  me  full  particulars  regarding  this  sta- 
tion. I  am  under  obligation  also  to  Mr.  David  White,  of  the  U.  S.  Geological 
Survey,  Washington,  D.  C.,  for  information  relating  to  the  geology. 

fBull.  Torr.  Bot.  Club,  Q:  350.     1879. 


—42— 

pit.  It  reappears  at  the  surface  nearly  two  miles  distant,  on  the 
level  flat  a  short  distance  from  the  river.  Here  it  is  called  the 
Blue  Spring.* 

As  nearly  as  can  be  ascertained  without  a  special  visit  to  the 
place,  the  formation  in  which  the  sink-hole  occurs  is  the  Bangor 
"rotten"  or  "Mountain"  limestone.  [See  note  III  in  thesis  by 
the  writer,  previously  mentioned. )  This  limestone  includes  a  few- 
beds  of  shale,  several  occurring  near  the  top,  so  that  the  presence 
of  the  clayey  habitat  is  easily  explained,  t 

SEVERAL  SUPPOSED  STATIONS. 

(a)  Chiapas,  Mexico.     In  his  Ferns  of  North  America,  J  Prof. 
Eaton  cites  the  Hart's-tongue  as  occurring  at  "Chiapas,  Mexico." 
The  specimens  (Eaton  Herbarium)  upon  which  this  determination 
was  apparently  based,  were  collected  in  ' '  cool  regions  ;  crevices  of 
rocks  in  the  bottoms  of  caves  (296-307)"  at  Chiapas,  Mexico,  by 
Ghiesbreght.  ||     They  are  not  to  be  referred  to  Phyllitis  Scolopen- 
dnum,  but  rather  to  Phyllitis  Linden!  (Hook. )  §  a  species  clearly 
distinct  from  the  former.     The  habitats  of  Ghiesbreght' s  and  Lin- 
den's specimens  are  rather  diverse,  but  not  more  so  than  those  of 
many  species  of  Polypodium  of  the  same  region. 

(b)  Sitka,  Alaska.     Milde^j  records  the  fern  from  "  Insula  Sit- 
cha,"  and  adds  "(herb.  caes.  Petrop.  horti  bot. ),"  indicating  that 
the  specimens  are  preserved  in  the  St.  Petersburg  Herbarium.     The 


*The  greater  portion  of  the  foregoing  has  been  kindly  contributed  by  Mr. 
nd  Mrs.  Joseph  H.  Lodge,  of  South  Pittsburg,  who  at  the  time  (1898)  of  their 
ivestigation  forwarded  several  living  plants  to  me.     I  have  described  the  sta- 
on  in  some  detail,  since  an  article  has  recently  appeared  (James  II.  Ferriss, 
Bull.  7:  98.  1899)  indicating  doubt  as  to  the  existence  of  the  fern  in  that 
tation  at  present.     It  is    evident  that   Mr.    Kerriss  missed  the   sink-hole  in 
question,  which  is  not  remarkable,  since  there  are  several  in  the  near  vicinity. 
tSafford  and  Killebrew,  Elem.  Geol.  Teiin.  p.  153. 
tD.  C.  Eaton,  Ferns  N.  Am.  \:  247.     1879. 

I  See  Cat.  Coll.  Ferns  So.  Mexico,  mainly  at  Chiapas,  by  A.  Ghiesbreght, 
1864-70.  This  pamphlet  (pp.  10),  kindly  loaned  me  by  Prof.  Underwood,  is 
evidently  a  reprint.  The  determination  of  the  ferns  was  accomplished  by 
Hall,  probably  under  Prof.  Eaton's  supervision.  The  two  sheets  have  been 
kindly  loaned  me  by  Prof.  A.  W.  Evans,  of  Yale  University. 

\ScolopendriumLindeni  Hook.,  well  figured  and  described  (Hooker,  Ic. 
PI.  II.  \:pl.  488.  1842)  from  specimens  collected  "on  old  oaks,  Chamulars,  Prov. 
Chiapas,  Mexico,"  by  Linden,  \\.  1543. 

,  Fil.  Europ.  et  Atlant.  90.     1867. 


—43— 

Hart's-tongue  is  not  uncommon  in  Japan,  and  may,  like  some 
other  species  (notably  Dryopteris  montana,  for  a  long  time  sup- 
posed not  to  occur  in  North  America),  have  an  eastern  extension 
across  into  Alaska,  and  down  the  Pacific  coast  into  British  Colum- 
bia. An  examination  of  the  material  in  the  St.  Petersburg  Herbar- 
ium, would  of  course  establish  the  identity  of  the  specimen,  but 
would  not  prove  the  authenticity  of  this  station,  since  it  has  fre- 
quently happened  that  plants  from  Russian  territory  upon  both 
sides  of  Behring  Sea  have  been  carelessly  and  indiscriminately 
labeled. 

(c)  I'ancoui'er  Island,  British  Columbia.     In  the  fall  of  1898, 
Mr.  Hamburg,  a  Swedish  botanist,  then  just  returned  from  a  col- 
lecting trip  in  the  West,  assured  me  that  he  had  recently  collected 
specimens  of  this  species  growing  upon  Vancouver  Island.     The 
statement  was,  unfortunately,  not  substantiated  by  specimens. 

(d)  Maniioulin  Island,  Canada.     Professor  W.  H.  Jenkins  has 
written  me  of  the  reported  occurrence  of  the  fern  upon  this  island 
in  Lake  Huron,  rather  more  than  a  hundred  miles  northwest  of 
Owen  Sound.     I  have  found  no  further  reference  to  its  occurrence 
here. 

(e)  Louisville,  Kentucky.     John  Williamson  *  was  inclined  to 
discredit  the  reported  station  near  Louz'sville,  mentioned  by  Dr. 
McMurtrie.  t    A   reference  to  Dr.   McMurtrie's  book  (Library  of 
Congress)  shows  the  following  entry  :  "  Asplenium  Scolopendrium 
Crt.     Hart's-tongue,"  along  with  a  few  other  ferns  reported  from 
the  vicinity.     It  may  be  taken  for  granted  that  the  Hart's-tongue 
was  not  confused  with  another  species,  as  the  chances  for  such  an 
error  are  indeed  small.     The  fact  that  Williamson  was  unable  to 
find  the  fern  might  indicate  either  that  it  has  disappeared  from  the 
vicinity,  that  -'vicinity"  was  made  to  include  a  considerable  terri- 
tory about  Louisville,  or  that  the  fern  has  since  been  overlooked 
near  the  city.     The  second  supposition  is  the  most  likely.     It  seems 
that  Williamson  and  later  botanists  would  have  found  the  fern  if  it 
still  occurs  in  the  immediate  vicinity  of  Louisville  ;  and  it  is  not 
probable  that    it    has  been    exterminated.     At    the   time   (1819) 


*  Williamson,  Ferns  of  Kentucky,  p.  iv.     1878. 

tH.  McMurtrie,  M.  D.,  Sketches  of  Louisville  and  its  Environs,  Ed.  i,  p.  229 
Louisvile,  1819.  This  book  includes  a  "  Florula  Louisvillensis,  or  a  catalogue 
of  nearly  400  genera  and  600  species  of  Plants,  that  grow  in  the  vicinity  of  the 
town,  exhibiting  their  Generic,  Specific,  and  Vulgar  English  names." 


—44— 

McMurtrie's  book  appeared,  the  Geddes,  N.  Y.  station,  was  the 
only  one  then  known  in  America,  having  been  recorded  by  Pursh 
just  five  years  previously. 

CONCLUSIONS. 

I  have  described  with  considerable  fullness  the  American  local- 
ities where  Phyllitis  Scolopendrium  is  known  to  have  been  found. 
The  striking  and  invariable  characteristic  of  its  environment  ap- 
pears to  be  an  affinity  for  limestone  rocks — the  Corniferous  in  cen- 
tral New  York  ;  the  Guelph  Dolomite  and  Clinton  in  Ontario ;  the 
Bangor  (Mountain)  limestone  in  Tennessee, — representing  consid- 
erable range  in  choice  of  formations.  The  question  naturally 
arises  :  Why  should  the  number  of  known  localities  be  so  small  ? 

As  a  matter  of  fact,  the  Hart's-tongue  is,  as  I  have  indicated, 
far  more  common  than  has  usually  been  thought ;  and  it  will  prob- 
ably turn  up  continually  in  some  of  the  less  explored  regions  of 
Canada,  especially  in  the  northwest.  It  appears  to  require  for  its 
best  growth  a  cool,  well-shaded  limestone  ravine,  talus,  or  sloping 
woodland,  with  rich  wood  soil,  for  the  most  part  sufficiently  porous 
to  allow  free  drainage,  but  firm  enough  to  retain  considerable 
moisture.  Such  conditions  occur  in  hundreds  of  glens  in  the 
United  States  and  Canada.  There  is  moreover,  quite  a  wide  vari- 
ance, in  the  wet  heavy  clay  of  the  South  Pittsburg  sink-hole,  from 
the  rich  porous  soils  and  loose  leaf  moulds  of  the  central  New  York 
stations,  or  the  scant  soil,  of  the  limestone  crevices  in  some  of  the 
Owen  Sound  stations.  The  South  Pittsburg  clayey  habitat  is,  to  be 
sure,  somewhat  anomalous,  but  it  appears  that  almost  any  soil 
upon  limestone  will  support  the  fern  under  the  right  temperature 
conditions,  perhaps  the  most  important  factor  now  operating  for 
or  against  the  fern's  survival  and  in  determining  its  future  distribu- 
tion, is  the  presence  of  a  constant  low  temperature.  The  James- 
ville  pit-lakes  have  a  uniformly  cool  temperature  from  day  to  day. 
The  Chittenango  gorge  is  deep  and  the  fern  is  well  shaded.  Ferns 
growing  from  fifty  to  seventy-five  feet  below  the  surface,  as  in 
the  irregular  South  Pittsburg  chasm,  cooled  by  a  waterfall,  can  be 
affected  only  in  slight  degree  by  extremes  of  temperature.  The 
Canadian  stations  are  mostly  near  streams.  It  is  true  also  that 
extremes  of  cold  do  not  seem  to  affect  this  fern  deleteriously.  It 
remains  evergreen  through  ordinarily  severe  winters,  except  when 
unduly  exposed  by  removal  of  protective  forest  growth.  And  so, 
while  the  fact  of  an  even  low  temperature  does  not  adequately 


—45— 

explain  the  causes  resulting  in  the  present  peculiar  distribution,  I 
think  it  does  throw  considerable  light  upon  the  relationship  between 
environment  and  present  distribution.  Since  arriving  at  this 
conclusion,  I  have  chanced  to  note  that  in  several  British  works, 
considerable  attention  is  paid  to  the  fact  that  the  Hart's-tongue 
occurs  regularly  "  in  caves,  on  the  seashore  and  in  other  cold  and 
damp  situations,  "  and  again,  "more  especially  about  the  mouths 
of  caves,  deserted  mines,  at  the  borders  of  wells,  where  there  is  a 
current  of  cold  or  moist  air." 

It  is,  of  course,  a  well  known  fact  that  plants  characteristic  of 
high  northern  latitudes,  are  found  in  more  or  less  abundance  upon 
mountain  peaks  of  the  more  temperate  regions  thousands  of  miles 
to  the  south.  The  accepted  explanation  is  :  that  during  the  glacial 
epoch  the  plants,  gradually  forced  south  by  the  advancing  ice  sheet, 
upon  the  northward  retreat  of  the  ice,  moved  up  the  mountains, 
seeking  to  maintain  accustomed  environmental  conditions,  *  or,  for 
the  same  reason,  advanced  to  the  northward.  Thus,  the  Hart's- 
tongue  occurs  in  America  mostly  in  the  north.  It  has,  as  I  have 
remarked,  been  usually  regarded  as  a  boreal  type.  I  believe  that 
it  was  once  far  more  common  than  at  present,  and  that  it  will  fre- 
quently be  found  in  the  north,  perhaps,  as  I  have  suggested, 
stretching  across  from  Asia  to  Alaska,  and  down  the  Pacific  coast. 
In  the  United  States  it  has  yet  to  be  seen  west  of  the  Mississippi, 
though  it  may  possibly  occur  along  the  upper  tier  of  States,  assum- 
ing that  it  follow  a  belt  parallel  to  the  lower  limit  of  glaciation. 
Especial  search  and  exploration  in  favorable  localities  would  not 
be  without  good  results  in  general,  and  would  very  likely  result  in 
further  extensions  of  range  for  what  has  commonly  been  regarded 
as  one  of  the  rarer  American  species  of  ferns. 


The  distribution  of  the  Hart's-tongue  in  Great  Britain  is  pecu- 
liar as  well,  and  has  been  commented  upon  by  Mr.  Druery  (Choice 
British  Ferns,  p.  14.  1888)  at  some  length.  He  remarks  that  its 
comparative  rarity  in  Scotland  is  the  more  unexpected  in  view  of 
the  "  innumerable  glens  which  abound  there  and  seem  a  very  beau 
ideal  of  a  habitat  for  it;"  and  adds,  moreover,  "this  fern  is  one  of  the 


*  All  especially  interesting  exposition  of  the  facts  and  causes  having  to  do 
with  the  distribution  of  species  (especially  North  American),  is  contained  in 
the  latter  portion  of  Dr.  Asa  Gray's  "  Memoir  on  the  Botany  of  Japan,  etc,"  in 
Mem.  Am.  Acad.  Arts  and  Sci.  II.  &'•  l859- 


-46- 

least  dainty  in  its  requirements,  seeming  to  have  no  antipathy  in 
the  matter  of  soil  or  position,  and  in  many  localities  thriving  in 
abundance  under  the  most  adverse  conditions."  Mr.  Druery  is 
inclined  to  the  belief  that  this  paucity  in  certain  sections  is  due  to 
the  "greater  or  less  predominance  of  certain  forms  of  minute  insect 
life.  *  *  Just  as  we  find  in  our  gardens  that  certain  vermin 
attack  and  destroy  certain  plants,  so  it  is  only  reasonable  to  as- 
sume that  either  the  spores  or  prothalli  of  these  ferns  are  the  favo- 
rite food  of  some  of  the  minuter  insects,  in  order  to  explain  the 
absence  of  adult  plants.  Climatal  conditions  are,  of  course,  a 
potent  factor,  but  do  not  account  for  all  the  phenomena  observed." 
Mr.  Druery  then  cites  the  cases  of  certain  exotic  ferns  which  attract 
slugs  or  snails,  and  must  be  grown  in  isolation.  Such  a  one  is 
Cainptosorus,  in  England.  "Other  plants,"  he  writes,  "are  espe- 
cially subject  to  the  attack  of  wood  lice  ;  and  so,  doubtless,  such 
special  appetites  characterize  also  the  minuter  and  microscopic 
insect  world,  and  as  it  is  manifest,  when  we  consider  the  myriads 
of  spores  which  are  shed  in  vain,  that  these  must  become  mainly 
the  food  of  such  tiny  creatures,  we  need  hardly  seek  further  for  a 
solution  of  the  mystery.  A  harder  or  softer  envelope  to  the  spore, 
or  a  more  or  less  attractive  flavor  in  this  plant  itself,  would  deter- 
mine for  or  against  its  survival  in  the  struggle  for  existence." 

It  seems  necessary  to  suppose  that  some  such  cause  operates  to 
effect  the  odd  distribution  both  in  England  and  America.  In 
March,  1898,  I  noticed  the  presence  of  one  of  the  Lace  bugs 
(Tingitidae),  upon  plants  at  Jamesville,  in  considerable  numbers. 
They  seemed  mostly  to  attack  the  spores,  but  were  found  appar- 
ently destroying  the  leaves  as  well.  Snails  are  often  found  depend- 
ing from  badly  eaten  fronds.  These  scant  observations  tend  to 
substantiate  Mr.  Druery' s  proposition.  It  is  indeed  perfectly  sup- 
posable  that  in  certain  of  its  various  stages  of  development,  it  may 
peculiarly  attract  vermin  to  itself,  which  so  greatly  impair  its 
vitality  as  to  lessen  its  chances  for  survival  and  reproduction.  A 
series  of  careful  observations  and  experiments  along  this  line  would 
be  of  the  greatest  interest. 

U.  S.  National  Museum,  \Yashington,  D.  C. 


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